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SMlTHSOiNIAN INSTITUTION 
UNITED STATES NATIONAL MUSEUM 



PROCEEDINGS 



OF niE 



UNITED STATES NATIONAL MUSEUM 



VOLUME 96 




UNITKD STA'n-.S 

GOVERNMENT PRINTING Of KICK 

WASHINGTON : lO^S 



ADVERTISEMENT 

The scientific publications of the National Museum include two se- 
ries, known, respectively, as Proceedings and Bulletin. 

The Proceedings, begun in 1878, are intended primarily as a medium 
for the publication of original papers, based on the collections of the 
National Museum, that set forth newly acquired facts in biology, an- 
thropology, and geology, with descriptions of new forms and revisions 
of limited groups. Copies of each paper, in pamphlet form, are dis- 
tribui-ed as published to libraries and scientific organizations and to 
specialists and others interested in the different subjects. 

The dates at which these separate papers are published are recorded 
in the table of contents of each of the volumes. 

The present volume is the ninety-sixth of this series. 

The Bulletin, the first of which was issued in 1875, consists of a 
series of separate publications comprising monographs of large zoologi- 
cal groups and other general systematic treatises (occasionally in 
several volumes), faunal Avorks, reports of expeditions, catalogs of 
type specimens, special collections, and other material of similar nature. 
The majority of the volumes are octavo in size, but a quarto size has 
been adopted in a few instances in which large plates were regarded 
as indispensable. In the Bulletin series appear volumes under the 
heading Contributions from the United States National Herharivmi, 
in octavo form, published by the National Museum since 1902, which 
contain papers relating to the botanical collections of the Museum. 

Alexander Wetmore, 
Secretary, Smithsoniam. Institution. 



^iPis 






II 



Cv 



CONTENTS 

Page 

BoHAHT, G. E. The phorid flies of Guam. No. 3205. Febru- 
ary 17, 11)47^ 397-416 

New genus: Parafannia. 

New si)ecies : Mcgasclia {.Ucgusdia) Hctifimur, J/, suis, M. 
stuntzi, M. parabasiscta, Chonoccphalus hirmitus, C. subglaber, 
PuUciphora wj/inani, P. nigriventru, Parafannia molluscovora. 

C'aldweix, John S. Neotropical lanternflies of the genus 
Phinctus in the United States National Museum, with de- 
scriptions of four new species. No. 3194. May IG, 1945 ^__ 177-184 
New siiecies : Phrirtus aordidufi, P. minulacanihis, P. punctatus, 
P. regalis. 

( "lakk, Austin H.. and Zetek, Jaaies. The onychophores of 
Panama and the Canal Zone. No. 3197. February 21, 
194G ^ 205-213 

CusHMAX, R. A. The ichneumon-flies of the genus Gryp- 
tanura Brulle. mainly tropical American. No. 3193. May 
23, 1945^ 139-170 

New genus: CremnocryptuH. 

New species: Cryptunura lubcrculdta, V. dicostuta, C. quadri- 
mncuUita, C. mrdiosirigosa, C. bicarinata, C. piccothoraoB, 
C. planiscutcllata, C. polUigaster, C. conica, C. ruficeps, C. 
scptctitrionulin, C. apophysis, C. biliiicata, C. rufa, V. lineati- 
fi'tiiiir, C. coxata, C. boUvicnsis, C. isthmus, V. cxcalibur, C. 
aviridccs, C grucilipcs, C. gracilis, C. gcnulis, (J. parancnsis, C. 
trnuiterebrata, V. mnnilifroiis. 

New combinations: Cryplnnuni mcxicana (Cres.son), C. uriza- 
bcnsis (Cameron), V. propinqua (Cresson), C. prciiosa (Vier- 
eck), V. vurirgnla (Hrulle), V. inccrta (('resson), Vrcmnocryp- 
tus spiiiifcrus (Cameron), C. cingulatus (Tosquiuet). 

. A generic revision of the ichneumon-flies of the 

tribe Ophionini. No. 3200. July 17, 1917' 417-482 

New gciK'ra : AUiji-h'iphiftu, Potophion, Sintoithinn, 'I'roplitipfiion, 
Clistorapha, Jforthonrura, Chilophion, Aulophion. 

New species: Auslralophinn iit/latus, Potophion cnudatus, Simo- 
phion aicnrinatuH, Trophophion tcniiicrps, liocthonvura arida, 
Htauropoctonus chczanus, Aulophion bicarinatus, A. vxcarinu- 

tU8. 
.New CDmliiiiatlDiis : l{!i i/iirlKijihion /I'luniiipt ntiis ( AsIiiik'JkI ) , J\'. 
ligulifrr (Morlcy), Aglnophion purpurnsn uh (Smilli), A. 
su)i\'ilriinum (Endt-rleln ), CliHtoruplin suhfuliginosa (Asli- 
nn'iMl). Ginophion costalis (<;r(;sson), Vliihtjihion iihnoruiis 
(Felt), Sliiuroii'>rt(,nii.H raricgaltts (Ucbldii). 



I)at<- of iiubllrntluii. 



IV PROCEEDINGS OF THE NATIONAL MUSEUM vol.86 

Page 

Davis, A. C. Review of the weevils of the tribe Ophryastini 

of America, north of Mexico. No. 3207. July 8, 1947 ^ 483-551 

New species: Eupagoderes griseus, E. aeneus, E. robustus, E. 

pilosus. 
New variety : Eupagoderes morUvallis approximatus. 

Fennah, R. G. New lanternflies (Fulgoroidea) from South 

America. No. 3189. May 9, 1945 ^ 95-104 

New genus : Iquitosa. 

New species : Pintalia quadrimaculata, P. marmorata, P. oMiqui- 
vitta, P. falcata, P. curvivitta, P. daedala, P. vomerifera, Iqui- 
tosa shannoni, Ateson seniiluteum, A. luteospersum. 

Fisher, Walter Kenrick. Echiuroid worms of the North 
Pacific Ocean. No. 3198. April 11, 1946 ^ 215-292 

New orders: Xenopneusta, Heteromyota. 

New genera : Bonclliopsis, Euhonellia, Nellobia, Lissomyema. 

New si)ecies: Thalassema steinhecki, Listriolobus pelodes, 
OcJietostoina octomyotutn, O. edax, Arhynchltc inamoenus, 
Bonelliopsis alaskana, Eubonellia valida, Nellobia eusoma, 
AcantJiohamingia parudola. 

New subspecies : Echiurus echiurus alaskanus. 

Fisher, W. S. New beetles of the family Eucnemididae from 
Central America and the West Indies. No. 3188. May 8, 

1945 1 79-93 

New genera : Neodiapodius, Hylotastella. 

New siiecies : Temnillus aspericollis, Neodiapodius buscki, Dro- 
maeolus pulcher, D. panamensis, Fornax poeyl, F. valerio, 
Plesiofornax nigrinus, Farsus convexus, ArrMpis cubanus, A. 
insularis, Dirhagus albofasciatus, Hylotastella schwarzi, Nema- 
todes exiguus. 

■ — . New cerambycid beetles belonging to the tribe 

Disteniini from Central and South America. No. 3201. 

November 26, 1946^ 329-333 

New species : Distenia lateralis, D. spinipennis, Cometes etnar- 
ginata, C. bicolor. 

Gahan, a. B. Eight new species of chalcid-flies of the genus 
Pseudaphycus Clausen, with a key to the species. No. 3200. 

November 22, 1946 ^ 311-327 

New species: Pseudaphycus mcritorius, P. abstrusus, P. malinus, 
P. anyustifrons, P. nmndus, P. Umatulus, P. meracus, P. alveo- 
latifrons. 

. Review of some chalcidoid genera related to 

Cerocephala Westwood. No. 3203. December 31, 1946 i__ 349-376 
New subfamily: Cerocephaliuae. 
New genus: Acerocephala. 



* Date of publication. 



CONTENTS V 

ruge 

Gahan, a. B. — Continued 

New species: Choctos;)U(i tabida, Accroccphala acnif/ma. 

New combinations: CIkx toKitila f rater (Girault), Th<ocoIax 
bakeri (Crawford), 'J heocolaxia viridinotnm (Dodd), T. insu- 
laris (Dodd), T. insuluris var. grandis (Dodd), T. perpuhhra 
(Dodd), 2'. pcrpulchra var. metaUica (Dodd), T. scoUjtivora 
(Ashiuead), T. pitijophthori (Aslnnead), Accroccphala atrovio- 
lacea (Crawford), Eupclinella canadouHis (Provancher). 

Gazin, C. Lewis. Machacroidcft cothen Matthew, the suber- 
tooth creodont of the Bi-idger Eocene. No. 3202. December 
16, 1946^ 335-347 

GujMore, Ciiarijjs W. Notes on recently mounted reptile 
fossil skeletons in the United States National Museum. No. 
3106. Fi'bruary 5, 1046^ 195-203 

. The osteoloory of the fossil turtle Testudo 

'pracextans Lambe, Avitli notes on other species of Testudo 
from the Oligocene of Wyoming. No. 3199. March 28, 
1946^ - 293-310 

GoijjMAN, E. A. (See under Kellogg, Remington.) 

Heinricii, Carl. The genus i^wTu/eZZaZeller: A contribution 
toward a revision of the American pyralidoid moths of the 

family Phycitidae. No. 3190. May 18, 1945^ 105-114 

New species : Funddln iiinohUia. 

Kellogg, Remixgtox, and Goldman, E. A. Review of the 

sjjider monkeys. No. 3186. November 2, 1944 ^ 1-45 

New .sui)si)ecies : Afcks gcoffroyi yucatanensis, A. g. panamensis. 

MiiJ.ER, Robert R. Hy porhamphiut patris, a new species of 
hemiramjihid fish from Sinaloa, Mexico, with an analysis of 
the generic characters of IhjporJiamphus and Ileniirani- 

phus. No. 3195. July 27, 1945^ 185-193 

New species: llyporluiinyltus jxitria. 

SciiULTZ, Leonard P. A revision of the American clingfishes, 
family Gobie.socidae, with descriptions of new geiioi-a and 
forms. No. 3187. December 30, 1944 ^ 47-77 

New gen«'ra : Acyrtun. InfratridniH, Afcon. 

New spe(!ies : Sicj/ascn hildchrandi. 

New sub.species : Cotylis niyri])ittnin woodxi. 

. A new genus and two new s[)ecies of i)erc()id 

fishes from New Guineji. family Centropomidae. No. 3191. 

March 6, 1945 ' 115-121 

New Bt'nu.s : Xcnamhu.iHia. 

New HfMi'cle.M : Xtndtnhassis honcHni, X. Himnni. 

' Date of puljllcatloii. 



VI PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

Page 

ScHULTz, Leonard P. Three new sciaenid fishes of the genus 
Ophioscion from the Atlantic coasts of Central and South 
America. No. 3192. April 25, 1945 ^ 123-137 

New species: Ophiosoion brasiUensis, 0. venezuelae, 0. pana- 
mensis. 

. A revision of the genera of mullets, fishes of the 



=■5 

family Mugilidae, with descriptions of three new genera. 

No. 3204. December 5, 1946 ^ 377-395 

New geuera : Xenomugil, Crenimugil, Heteroitiugil. 

Zetek, James. (See under Ci.ark, Austin H.) 



^ Date of publication 



ILLUSTRATIONS 

PLATES 

Follow I iig 
page 

1. Lower side of head of Votylis microi^pUus (ri.wlcr) and of (\ 

nigripinnis niyripinnifi I'oters 64 

2. New South Aiuoiicau FiilKoroidea 104 

3. South American Fulgoroidea 104 

4-6. The pMius FundcUa 114 

7. Cephalic process in Phrictus 180 

8. Valvula and geuitalia in Phrivttis 180 

0,10. Species of Phrictus: Dorsal aspect 180 

IL Hyporhamphua patris, new species 193 

12. Mounteil skeleton of Crocodiliis clavis Cope 203 

l.S. Skull and k)wer jaw of Crorodilus clavis Cope 203 

1 1, ir». Skull of Crocodiliis clavis Cope 203 

16. I»isea.sed dorsal vertebrae of Crocodilns sp 203 

17. Tail, pelvis, hind linihs, and feet of Corythnsaurus casuarius Brown_ 203 

18. Skin impre.ssions of Conjthosnunis casuarius Brown 203 

19. Restoration of Cori/thosaurus casuarius Brown 203 

20. IJchiurus cchiurus alaskanus, new subspecies 292 

21. Listriolobus pelodes, new species, and Ochetostoma octomyotum, 

new species 292 

22. Listriolobus pelodes, new species 292 

23, 24. Ochetostoma octomyotum, new species 292 

2r». Arhynchite iuamoetius, new species 292 

26, 27. IloneUiopsis aUiskatia, new genus and species 292 

2S. Euhonellia ralida, new frenus and species 292 

20, ;}0. Xellohid cusoma, new genus and species 292 

;{1,32. Acunttiohaminf/ia paradola, new species 292 

^'.i-?,~). Frechis enupo Fislier and MacClinitie 292 

1^6. Urechis caupo, F. chilcnsis, and U. unicinctua 292 

37. Frcchis caupo and comniensals 292 

38-41. Tcstudo praeextans Lanibe 310 

42,43. Testudo laticunea Cope 310 

44. Testudo f/uadrala Cope; Testudo praeextans Lambe 310 

45-46. Mnrhnrroidrs cothen Matthew 347 

47. Si>f'cies of Choetospila, Thcocolax, Cerocephala, Acerocephala, and 

Theocolaxia 376 

48. Species of Thcocolax, Cerocephala, and Acerocephala 376 

40-.''>6. Ichrienmon-flios of the tribe Ophionlnl 482 

Ti:XT FICTIRES 

PnKo 

1. Distribution of the forms of spidernionkeys (Atcles) In South Amer- 

ica (map) 12 

2. Distribution of the forms of spider monkeys (Ateles) in Middle 

America (map) 14 

3. Holotype of Xmnmhassis honcHsi, new species 119 

4. IIolotyi)e of Xctuimbassis simoni, new speci»'H 121 

f). Diagranunatic sketches of snout and lower Jnw of Bairdiella chrys- 

ura fLacep<>de), Stellifer rastrifer (Jordan), and Ophiosdon 

typicus Cill 12.'» 

6. ITolotyi»e f»f Ophi'iscion hrasilicnsis, new species 129 

7. Iliilotyjx' of (tjiliioscitni rcnczurhic, new species 132 

8. JIololyp<' of Ojihi'iscioti paunmcnsis, new species 13.'> 

9. Sketch of hearl regions of Ifyporhamphus and Ifcmiramphwt 180 

VII 



VIII PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

Page 

10. Lissornyema mellita (Couu) 223 

11. Thalassema steinbecki, new species 231 

12. Ldstrioloius pelodes, new species 235 

13. Listriolohus pelodes, new species 237 

14. Ochetostoma edax, new species 244 

15. BonelUa viridis : 251 

16. BonelUopsis alaskana, new species 252 

17. Urechis caupo Fisher and MacGiuitie 266 

18. Urechis caupo Fisher and MacGinitie 267 

19. Urechis caupo Fisher and MacGinitie 273 

20. Skull of Testudo pracextans Lambe 294 

21. Skull and lower jaw of Testudo praeextans Lambe 295 

22. Palatal view of the skull of Testudo praeextans Lambe 296 

23. Lower jaw of Testudo praeextans Lambe 297 

24. Carapace of Testudo praeextans Lambe 298 

25. Plastron of Testudo pi-aeextans Lambe 303 

26,27. Pelvis of Testudo praeextans Lambe 304 

28. Sketch of the maxillary, premaxillary, and preorbital bones of 

Mugil cephalus 388 

29. Sketches of the maxillary, premaxillary, and preorbital bones of 

Myxus elonfjatus 390 

30. Sketches of the maxillary, premaxillary, and preorbital bones of 

Chelon cheio 390 

31. Sketches of the maxillary, premaxillary, and preorbital bones of 

Trachystoma petardi 393 

32. Diagram of the possible I'elationships of genera of the Mugilidae 395 

33-35. Phoridae of Guam 400 

36-38. Phoridae of Guam 401 

39-41. Phoridae of Guam 404 

42-45. Phoridae of Guam 405 

46-48. Phoridae of Guam 408 

49. Genital structures of Eupagoderes cf. deserttis-caUfornicus sec- 

tion 486 

50. Sapotes puncticollis Casey 489 

51. Eupagoderes geminatus Horn 497 

52. Eupagoderes griseus, new species 499 

53. Eupadogeres aeneus, new species 500 

54. Eupagoderes mortivallis Fall 502 

55. Eupagoderes rohustus, new species 504 

56. Eupagoderes decipiens (LeCoute) 505 

57. Eupagoderes spcciosus (LeConte) 510 

58. Eupagoderes niarmoratus Fall 512 

59. Eupagoderes sordidus (LeOonte) 514 

60. Eupagoderes simulans Van Dyke 515 

61. Eupagoderes argentatus (LeConte) 516 

62. Eupagoderes lucanus Horn 518 

63. Amydrogmus variabilis Pierce 521 

64. Ophryastes tvickhami Sharp 523 

65. Ophryastes latirostris LeConte 526 

66. Ophryastes symmetricus Fall 528 

67. Ophryastes sulcipennis Casey 529 

68. Ophryastes sulcirostris (Say) 530 

69. Ophryastes porosiis LeConte 532 

70. Ophryastes tuberosus LeConte 534 

71. Ophryastes ovipennis Sharp 535 

72. Ophryastes vittatus (Say) 537 

73. Tosastes coarctatus Champion 540 

74. Tosastes globularis Pierce 542 

75. Tosastes ovalis Pierce 544 

76. Tosastes cinerascens Pierce 546 

77. Rhigopsis effracta LeConte 548 



PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 




issued n^t''t\»>L^JH*j;l ^V <^< 



SMITHSONIAN INSTITUTION 

U. S. NATIONAL MUSEUM 



Vol. 96 WMhington: 1944 No. 3186 



REVIEW OF THE SPIDER MONKEYS 



By Remington Kellogg and E. A. Goldman 



Field studies have shown that the forest-inhabiting spider monkeys 
of the New World are «nl)jpct to ])lasmodial infections under nat- 
ural conditions and consequently, since they are readily obtainable. 
are now being used in laboratory studies of the malaria Plasmodium. 
Spider monkeys are readily tamed and very adaptable to a life in 
captivity. Owing to their d(jcility they are easily handled by labo- 
ratory assistants, and it is not difficult to provide suitable vegetable 
fofxl for them. For these reasons it is important to establish, if pos- 
sible, the valid geographic races of these monkeys, to determine in so 
far as available collections permit the geographic ranges of the recog- 
nized forms, and to set forth tlie characteristics by which the several 
races can be recognized with s(jme degree of certainty. 

The spider monkeys form a compact group, or subfamily, the 
Ateliiiae, of the primate family Cebidae, 'I'liey are characterized by 
slender body, very long, slender limbs, and long tail, naked beneatli 
distally and prehensile; the hand has only four functional fingers, the 
thumb being usually vestigial or absent. 

These monkeys, in structure and hai)its, sliow higli specialization 
and adaptation to a strictly arboreal life. They arc limited in range 
to tlie unbroken tropical ff)rests from southern Mexico to northern 
Matto Gnjsso, Brazil, and central Bolivia in South America. In the 
forest areas inhabited, oflen including steep mountain slopes, largo 
trees have interlocking branches through which the spider monkeys 
are able to travel in long leaps at amazing speed, using the tail con- 
stantly as a powerful grasping organ for balancing, and swinging to 
differing levels. A spider monkey in a cage is seen at a disadvantage 



2 PROCEEDINGS OF THE NATIONAL MUSEUM vol. m 

compared, for example, with so active an animal as the gibbon, but 
the reason for this seems to be that room is lacking for the exercise of 
its full powers. It is interesting to speculate on the comparative 
speed that might be attained under natural conditions by these widely 
different but both outstandingly active New World and Old World 
representatives, respectively, of the order to which they belong. In 
following spider monkeys in the forest a man may keep up by run- 
ning if the ground is clear, but if impeded by having to go around 
underbrush he is quickly left behind. Spider monkeys are considered 
very good to eat by the native populations of many countries and 
are, therefore, in danger of extermination. 

Many names for spider monkeys have been based on inadequate de- 
scriptions of animals from unknown localities, and if a type specimen 
was used it may no longer be extant. The result has been great con- 
fusion in the literature bearing on the identification of material, and 
of the distribution of the nominal species recognized little has been 
known. 

The status even of the generic name Ateles has been open to ques- 
tion. The spider monkey S[imia\ Sapajus paniscus was included 
among the 14 monkeys allocated to Sapajus when this subgenus was 
proposed by Kerr (The Animal Kingdom, CI. I, Mammalia, p. 76, 
1792). No genotype seems to have been designated for the subgenus 
Sapajus Kerr, unless the statement made by J. A. Allen (Bull. Amer. 
Mus. Nat. Hist., vol. 7, p. 181, June 20, 1895) that Sapajios of Kerr is 
equivalent to Cebus Erxleben, 1777, can be interpreted as restricting 
the application of the term Sapajus to the species of Gehus included 
among the 14 monkeys mentioned above. In order to eliminate any 
possible misinterpretation, the genotype of Sapajus Kerr is here desig- 
nated as S\i7nia'\ Sapajus capuclnus Kerr {op. cit., p. 78, 1792), which 
is the same as S[i77iia'] capiwina Linnaeus., 1766 {nee 1758). Inasmuch 
as the genotype of Gehus Erxleben, 1777, has been fixed (Elliott, Field 
Columbian Mus. Publ. 115, zool. ser., vol. 8, p. 560, Mar. 4, 1907) by 
subsequent designation as Sitnia capucina Linnaeus, 1766 {nee 1758), 
which in turn is equivalent to Gehus nignvittatus Wagner (see 
Cabrera, Rev. Soc. Argentina Cienc. Nat., vol. 16, pp. 21-22, 1939), 
Sapajus Kerr is herewith relegated to the synonymy of Gehus Erxle- 
ben. 

Neither opinion 147 (On the principles to be observed in interpreting 
article 34 of the International Code in relation to the rejection, as homo- 
nyms of generic and subgeneric names of the same origin and meaning 
as names previously published. Opinions and declarations rendered 
by the International Commission on Zoological Nomenclature, London, 
vol. 2, pt. 14, pp. 123-132, Sept. 30, 1943) nor article 34 (International 
rules of zoological nomenclature, Proc. Biol. Soc. Washington, vol. 
39, p. 86, July 30, 1926) contains an express ruling as to whether a 



THE SPIDER MONKEYS — KELLOGG AND GOLDMAN 3 

ijiven oreneric nainc is a hoinonyni oi' anolluT piwiously jmblislied 
ofenerio name that has the same ori<2:iii and meaning but that dill'ers 
from the hitter in more than one letter. If the "one letter" rule is 
strictly adhered to, then Sopajou Lacepi'de (Tableau des divisions, 
sous-divisions, ordres et genres des mannniferes, p. 4. Published as 
supplement to Discours d'ouverture et de cloture du cours d'histoire 
naturellc, et tableaux nu'thodiques des mannniferes et des oiseaux, 1799) 
is the oldest available name for the spider monkeys. The genotype of 
Sapajou Lacepede is Sapaj&u paniscus Lacepede=/S'/wMf paniscus Lin- 
naeus. Pending an opinion from the International Commission on 
Zoological Xomenclature, the generic name Ateles E. Geoffroy (Ann. 
Mus. Hist. Nat. Paris, vol. 7, p. 2G2, 1806) , of which the genotype is 
likewise Simla paiiiscus Linnaeus, is here retained for the spider 
monkeys. 

We have been unable to reach any satisfactory conclusions regarding 
the identity of one of the descril)ed spider monkeys allocated to this 
genus. The '"Antigua Monkey" was described by Pennant (His- 
tory of quadrupeds, vol. 1, p. 206 (no. 212), 1781) as follows : 

Mfonkey] with a short nose; black face, hair on each side long; back and sides 
orango and black, intimately mixed ; holly white; outside of lops black ; inside ash- 
I'olorotl ; tail of a dusky ash; its length 20 inches; that of body oightocn. 

Lately in possossion of Richard Morris, Esq. of the Navy-Offico; brought from 
Antigua; but its native place uncertain; very good-natured, lively, and full of 
tricks; frequently hung by its tail. 

This monkey subsecpiently became the basis for [^Sinua] Sapajus 
variefjatuH Kerr (The Animal Kingdom, p. 79, 1792) and Simia antiqu- 
ana Pechstein (Thomas Pennant's Allgemeine Uebersicht der vierfiis- 
sigen Thiere, vol. 1, p. 227, footnote, 1799). In our judgment these 
names apply clearly to a spider monkey, and quite likely to one of the 
Middle American subspecies, but since the characters given are not 
sufficient to identify the race positively, we believe both shoidd be held 
unident ifiable. '^\n<ni Atclcs variegatws (Kerr) antedates .1 tchfi va/ic 
fjahiJi ^^'agner, the latter being preoccupied must be supplanted by 
Ateles hehehutli E. Geoffroy, which also has priority. 

Despite strongly contrasting patterns of color, dilfering lengths of 
pelage, and to some extent varying tail and foot proportions, the strik- 
ing similarity of all the spider monkeys in the more essential cranial 
features seems to indicate close reiationsiiips. The nominal sjxries 
arc here reduced to foiii-. mimI it seems to us probable that additional 
study of the genus based on moiv complete collections may result in a 
further le.ssening of the number. The most divergent forms are the 
I'lack paniscus of French Ciuiana and the silvery-bellied yucatanensis 
(jf Qiiintana Koo. Perhaps the most clearly defined line of demarca- 
tion between species, as we understand them, is in eastern Panama, 
where the range of the deep reddish panamcnsM, a member of the 



4 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

geoffroyi group, meets or closely approaches the range of the nearly all 
black Tohustus. The Rio Amazonas, however, seems to be an effective 
barrier separating the ranges of A. paniscus paniscus and A. helzebuth 
marginatiis. 

Several large series of specimens from the same locality have 
afforded us much information concerning the range of individual 
variation that may be expected in the genus Ateles. The thumb is 
normally represented by a short metacarpal, which ordinarily is not 
visible externally. When, in occasional individuals, a short proximal 
phalanx is retained it constitutes the vestigial nailless thumb. In 
two specimens of velleroms (U. S. N. M. Nos. 74662, 74663) from 
Santa Efigenia, Oaxaca, and in one of the same form (A. M. N. H. 
IS'o. 123282) from Cantoral, Honduras, these thumbs are present on 
one hand and not on the other. In one of pananiensis (A. M. N. H. 
No. 141980) from Canas Gordas, Costa Rica, the thumbs are present 
on both hands. In a specimen of hyhridus from Guaimaral, Colom- 
bia, vestigial thumbs on both hands bear small nails. Lonnberg 
(Arkiv for Zool., vol. 32A, No. 25, p. 8, July 18, 1940) describes a 
well-developed thumb with a well-developed nail on one hand only 
in a specimen of paniscus^ a small thumb present on the other hand 
being nailless. All three specimens of paniscus (A. M. N. H. Nos. 
94134, 94135, 94136) from Rio Jamunda, near Faro, on the north bank 
of the Rio Amazonas, have thumbs on both hands, as already re- 
corded by Tate (Bull. Amer. Mus. Nat. Hist., vol. 76, p. 215, Oct. 20, 
1939). Two others from Lago Cuipeua, farther east on the same 
bank of this river, however, are thumbless. Lonnberg (op. cit.) 
refers to the absence of thumbs in specimens of Ateles ater 
[= chameh] from various localities, but they are present on both 
hands in a skin (C. M. No. 2775) from Rio Yapacani, Santa Cruz, 
Bolivia. The evidence thus presented indicates that vestigial thumbs 
occur irregularly throughout the genus, and no specific significance 
is attached to their presence or absence. 

Certain cranial details believed by some authors to represent dis- 
tinctive characters prove to be too variable to be of much value. As 
Lonnberg [op. cit.) has pointed out, some of the features mentioned 
as characters by Tate {op. cit.) in comparing species are subject to 
a wide range of variation. The antorbital or "malar foramen" varies 
from one or two "pin-holes" to rounded openings 5 mm. in diameter 
through the jugal. These often differ in size and number on the 
two sides of the same skull. Tate also refers to "small, triangular 
pterygoids without pointed tips" as a cranial character in paniscus. 
In the Cebidae the fossae between the true pterygoids and their ex- 
ternal reduplications are large and deep in Cehus., reduced to a ves- 
tige in Alouatta., and entirely wanting in Ateles. The external 
reduplications of the pterygoids have the tips broken off in the 



THE SPIDER MUNKEYS KELLOGG AND GOLDMAX 5 

spei-inieiis of paiif.fcus exiuninotl by Tate, but tlu'^^e wiii^like thin 
plates are in reality lar<ier and more extended posteriorly in these 
and in other specimens of pan>,'<ciis from (he Amazonian refrion than 
usual in this genus. Other characters that have been ascribed by 
Tate to paniscus but that have not been found to be constant are the 
small size of post<ilenoid jjrocesses, unexpanded zy^jfoniata of nearly 
uniform depth, and narrow teeth. The maxillary tooth rows in the 
^^enus as a whole are normally straight and convergent anteriorly, 
although skulls with parallel or arcuate tooth rows are less frequently 
observed. All the teeth in the maxillary tooth rows may be large 
or small, and the individual teeth vary in size in relation to one 
another. The posterior molars appear to be the most variable. 
They are usually similar to the first premolars in crown area but 
may be either larger or smaller. Examination of a large number of 
spider-monkey skulls shows that the cranial and dental features men- 
tioned are subject everywhere to about the same wide range of indi- 
vidual variation. Schultz (Journ. Mammalogy, vol. 7, No. 4, pp. 
28G-305, Nov. 1926) has discussed the relative ranges of variation in 
skull measurements and in the formation of the forehead in a series 
of Atelen taken in the immediate vicinity of one camp on the bank 
of the Rio Yoya in Nicaragua. 

Beyond the general similarity and the range of the usual individual 
variation in the forms of Atrles there are, however, a few cranial 
features that seem to be worthy of consideration. All the races of 
Ateles geoffroyi agree closely in the more important cranial details, 
but in the more northern subspecies vellerosus and yueatanens-ia the 
brain case is slightl}' narrower than in panamensis and other Middle 
American races. The South American species A. paniscv-f and ^1. 
hrlzehiith have a longer rostrum, a more strongly inclined backward 
facial piofile, and moie elongated nasal openings than geoffroyi. 
In j)anisf'Uf<, which api)ears to be the most divergent in cranial de- 
tails, the pf)stei-ior upi)er molars scarcely reach the transverse plane 
of the anterior roots of the zygomata, while in geoffroyi they nonnally 
jjass well beyond this plane; other j)ecidiar chaiaeters in pcmisms 
are the broad, somewhat swollen base of the rostrum and the poste- 
rior exten.sion of the external reduplications of the pterygoids. In 
cranial details, as well as in geogi-a|)hic position, At< Us fii.sc/a ps ru- 
bu.shiM is somewhat intermediate iM'tween geoffroyi wwd jxniiftcus. 

Specinu'ns examined in series have shown tliat altliough sjnder 
monkeys present considerable variation in color the vaiiation is 
within limits beyond which it does not normally pass, except as the 
usual intergradation that may be expected Ix-tween regional races or 
subspecies. The coloration, length of ptdage, and othei* external 
features, when determined by examination of specimens in suflicient 
numbers, prove to be more reliable indices to subspecific relationship 



6 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

than any cranial characters found by us. In Middle America, for 
example, skulls of vellerosus from Mexico are distinctly narrower, 
with less noticeable widening of the brain case, than those of fana- 
me^isis, but differ little in other respects and despite great differences 
in color are obviously assignable to the same species. Unlike many 
other primates the sexes in the spider monkeys differ little, if at all, 
in size. One of the largest skulls of geoffroyi is that of an old female 
from Lavala, Nicaragua. 

For loan of pertinent material our thanks are tendered to the fol- 
lowing: Dr. Thomas Barbour, director, and Barbara Lawrence, Mu- 
seum of Comparative Zoology, Cambridge, Mass.; Dr. Adolph H. 
Schultz, Laboratory of Physical Anthropology, Johns Hopkins 
Medical School, Baltimore, Md. ; Dr. H. E. Anthony and George G. 
Goodwin, American Museum of Natural History, New York, N. Y. ; 
J. Kenneth Doutt, Carnegie Museum, Pittsburgh, Pa.; Dr. H. C. 
Oberholser, Cleveland Museum of Natural History, Cleveland, Ohio; 
Dr. William H. Burt, University of Michigan, Museum of Zoology, 
Ann Arbor, Mich., and to the latter also for the opportunity to 
examine specimens in the Donald R. Dickey collection. 

In lists of specimens examined, the following abbreviations are 
employed : 

A. H. S., Collection of Dr. Adolph H. Scbnltz. 

A. M. N. H., American Musenm of Natural History. 

C. D., Collection of Donald R. Dickey. 

C. M., Carnegie Museum. 

C. M. N. H., Cleveland Museum of Natural History. 

M. C. Z., Museum of Comparative Zoology. 

U. M. M. Z., University of Michigan, Museum of Zoology. 

U. S. N. M., United States National Museum. 

All measurements are in millimeters. The external measurements 
were taken in the flesh by the collector as follows : Total lengthy nose 
to end of terminal vertebra; tail, upper base of tail to end of terminal 
vertebra; hind foot, back of heel to end of longest nail on digit. The 
following cranial measurements of typical adults, unless otherwise 
stated, were taken with vernier calipers by the authors: Greatest 
length, distance from anterior tip of premaxillae to inion or extreme 
posterior median point of brain case ; orhital width, distance between 
outer margins of orbits; 'postorbital constriction, least width at con- 
striction behind orbits; width of hrain case, greatest width of brain 
case at or over mastoids ; zygomatic hreadth, greatest distance between 
outside surfaces of zygomata ; inaxiUary tooth row, distance from front 
of canine to back of posterior upper molar at alveolar borders. 

The following treatment is based on a review of the scattered lit- 
erature and an examination of 251 specimens representing all the 
recognized species. The work has been done at the request of the 
Board for the Coordination of Malarial Studies of the National Re- 



THE SPIDER MONKEYS KELLOGG AND GOLDMAX 7 

search Council in the hiboiiitorics of the Division of Miimnials luid of 
the V. S. Fish and Wildlife Service in the United States National 
Museum. The nuips were drawn by Mrs. Kiithoryne C. Tubb, of the 
Fis-h and Wildlife Service. This review is inteniled to afford a work- 
ing knowledge of the genus, pending more exhaustive revision as the 
species become better known. 

KEY TO SPIDER MONKEYS (GENUS ATELES) ' 

A. Gi'iieral coloration of entire back ehieflj' black. 

B. Back and belly, a.s well as lin/bs ami tail, wbolly deep glossy black. 

C. Hairs on bead wholly black (except for short, white mustache whiskers 

on upi)er lip and sparse, short, white chin whiskers) ; pelage deep glossy 

black. 

D. Foot rather larj^e (length, hi^l to end of longest toe, 190-220 mm.). 

E. Tail normally bushy in adult (especially on upper ba.sal half, longest 

hail's on upperside .50-100 nun. in length) ; face normally flesh color, 

occasionally freckled or black; hairs on back silky, lax, and long 

(majority of hairs on midline of back 75-150 mm. in length) ; tail 

often more than twice length of head and body. (North bank of 

Amazon east of Manfios and north to Caribbean coast of Guianas. ) 

paniscus 
EE. Tail much less bushy but more densely furred in adult (longest 
hairs on upi>erside 4(M');j mm. in length); face wholly black; 
hairs on back .shorter (majority of hairs on midline of back 
75 nrm. or less) ; tail often more than twice length of head and 
bod J'. (Western Mat to Gros.so, eastern Bolivia, and north- 
eastern Peru to Rio Solimoes and Rio Jurua, Brazil.) chamek 

DD. Foot smaller (length, heel to end longest toe, 100-170 mm.) ; tail not 
noticeably bu.shy (the longest hairs on upperside 30-50 mm. in 
length) ; face wholly black, except for short, white nuistache 
whiskers on upper lip and sparse, short, white chin whiskers; 
hairs on back harsh and of medium length (majority of hairs on 
midline of back 40-70 mm. in length) ; tail variable in length, 
occasionally nearly twice as long as head and body. (Serranfa del 

Darien, Panama, to .southwestern Colombia.) robustus 

CC. Hairs on head not wholly black; semilunar forehead patch and side 
whi.skers on face white; remainder of body, lind)S, and tail deep 
glo.ssy black; foot nre<lium (length, heel to end longest toe, 170-185 
mm.) ; tail vari;d»le in length, occasionally more than twice as long 
as head and body. (Slate of Parfi. Brazil, .south of Amazon, between 

Rios TajiajTiz and Tocatitlns. ) marginatus 

JiB. liack and belly, as well as limbs and tall, not wholly deep glossy black; 
hairs on back black at hnsi', and either black dislaliy or with lii>s 
tinged with either burnt umlter or seal brown. 
F. Crown cap tawny-oUve, yellowish wood brown, oi- willi black hairs lipped 
with burnt umber; tuiderparts similar in color to \ipjH'riiarts; entire 
body, Including limb.s and tail, covered with black hairs tinged dislaliy 
with burnt umber; foot small (leiiglli. heel to end longest toe, l.'.O-170 
mm.) ; tail variable in length, usually one-flfth or more longer lh:in 
head and body. (Pacific side of cordlllera of Ecuador.) fusciceps 



' ('..lors niffitlono*! In thU kpy are nhown In Ilotjort RIdKwny'fl 'Tolor SlandnrdH nnd 
Color Nomenclature," 4.3 pp.. r.3 pig. WuHliinKton, 1012. 



8 PROCEEDINGS OF THE NATIONAL MUSEUM vol.98 

FF. Crown cap black, not tawny-olive, yellowish wood brown, or with hairs 
tipped with burnt umber ; underparts conspicuously lighter in color 
than upperparts, line of demarcation between upperparts and under- 
parts being sharply defined ; general coloration of crown cap on head 
as well as rest of upperparts black ; in some individuals these black 
hairs under strong light having a seal-brown tinge ; triangular fore- 
head patch yellow, golden, brownish yellow, or white (occasionally 
either entirely absent or more or less hidden by anterior stiff black 
hairs) ; side whiskers on face, when present, whitish or huffy (not 
present in some specirtfens) ; outer surfaces of forelimbs to elbows 
black ; outer surfaces of forearms to wrists generally blackish but 
occasionally similar to underparts and intermixed with long dusky 
overhairs ; outer surfaces of hind limbs usually blackish to knees or 
below, but in some specimens (particularly in eastern part of range) 
outer surfaces of hind limbs, including thighs and region around base 
of tail, distinctly straw-colored in ground color overlaid with sparse 
long black hairs (not numerous enough to affect general ground color) ; 
hairs on throat normally black, sometimes grayish, more or less 
concealed by black hairs ; remainder of underparts, inner surfaces 
of fore and hind limbs, as well as under surface of tail, yellow, 
cinnamon-buff, dull cream-buff, or pale olive-buff; tail normally 
bicolored, upper surface being black and under surface similar to 
that of underparts (in occasional specinfens upper surface of tail 
approximately same as that of under surface but more heavily inter- 
spersed with long black overhairs) ; foot large (length, heel to end 
longest toe, 190-200 mm.) ; tail variable in length but often almost 
twice length of head and body. (From near junction of Rio Caura 
with Rio Orinoco in Venezuela south to Rio Negro in Brazil, westward 
to Colombia east of Cordillera Oriental (Mambita), Ecuador east of 

Andes, and northeastern Peru.) belzebuth. 

AA. General coloration of entire back not chiefly (jet) black. 

O. A conspicuous white triangular forehead patch ; general coloration of i:pper- 
parts wood brown, darker on head and upper back and lighter on lumbar 
region and thighs ; underparts, inner surfaces of fore and hind limbs, and 
under surface of tail whitish or buffy ; side whiskers on face variable, 
sometimes whitish or buffy and sometimes dark wood brown. (Magda- 

lena River Valley and adjacent mountains, Colombia.) hybridus 

GO. No conspicuous white triangular forehead patch ; general coloration of 
upperparts not wood brown. 
H. General coloration of upperparts, except for head and shoulders on some 
individuals, tending toward light mahogany red ; underside of tail, at 
least on basal half, deep cinnamon-rufous to ferruginous ; back and 
shoulders grading from dark rusty reddish to ferruginous or burnt 
sienna, with varying admixture of overlying blackish hairs ; underparts 
similar in color to back but with less noticeable admixture of blackish 
hairs ; face, crown cap on head, and median streak on back of neck 
blackish ; sides of neck covered with a mixture of pinkish-bufE, cin- 
namon-buff, and blackish hairs ; outer surfaces of fore and hind limbs 
either black or blackish to knees and elbows ; outer surfaces of limbs 
below knees and elbows either black or with varying admixture of black 
and ferruginous hairs ; a distinct ferruginous or cinnamon-buff streak 
along inner surface of arm from armpit to elbow. (Western side of 



THE SPIDER MONKEYS — KELLOGG AND GOLDMAN 9 

central curdillera of Costa Uiea (lilo I'iiris) south to Cordillera de 

San Bias (Cerro Azul) of eastern rauama.) panamensis 

////. Cieneral coloration of upporparts, except for head and shoulders, not uni- 
form light mahogany red ; underside of tail not deep ferruginous. 
/. GeruMal coloration of uppt'riiarls as well as underparts light hufC over- 
laid with dusky tipiK-d hairs; side whiskers on face silky light buff; 
dark markings on head and limbs usually inconspicuous; crown cap 
on head grading from black to buff tinged with brownish ; stiff long 
black hairs above eyes concealing to a variable extent the white or 
buff triangular forehead patch ; elbows and outer sides of forearms 
more or less distinctly blackish ; elongate black patches on knees, the 
hairs black to roots. (Southern Nicaragua, Greytowu to Lake Mana- 
gua.) I . geoffroyi 

//. General coloration of upperparts as well as underparts not light buff. 
J. Chest and belly normally near tawny or cinnamon-rufous. 

K. General coloration of upperparts golden yellowish to cinnamon- 
rufous in tone but obscured by numerous overlying black-tipped 
hairs; crown cap black, the hairs black to base; underparts and 
flanks near tawny or cinnamon-rufous ; tail black above, more or 
less mixed with tawny below from base to near callosity; side 
whiskers on face blackish. (Eastern side of central cordillora 

of Costa Rica. ) ornatus 

KK. General coloration of upperparts lighter or duller than preceding. 
L. General coloration of upperparts, except for head and shoulders, 
clear tawny, intermixed with black-tipped hairs ; sides, thighs, 
and belly paler, tawny or cinnamon-buff, with somewhat paler 
tone extending downward to wrists and ankles on inner sur- 
faces of fore and hind limbs; shoulders and outer surfaces of 
fore and hind limbs black; tail .sharply bicolored, black above 
to tip, and tawny below; crown cap either black or dusky (in- 
dividual hairs buff or cinnamon-buff at base) ; side whiskers on 
face cream color. (Azuero Peninsula, Panama. )_-azuerensis 
LL. General coloration of upperparts, except for head and shoulders 
in some individuals, buckthorn brown to Mars brown, sparsely 
or Jioticeably intermixed with blackish hairs (especially on 
midline of back in sonit? individuals) ; belly honey yellow to 
tawny, «'Xten(liiig downward on inner surfaces r>f hind limbs to 
ankles, and a lighter tone extending downward on inner sur- 
faces of forellmbs to wrists; outer surfaces of fore and hind 
limbs usually black (in some individuals restricted more or 
le8.s to elbow and knee patclies) ; tail usimlly sharply bicolored, 
bhnk or diisky above to tip and tawny below (when otherwise, 
mixed light and dark hairs cover tall above and below) ; an- 
terior part of crown jiatch more or less suffused with cliuia- 
nion buff, owing to liglil basal color of liairs and tending to 
form a transverse band across forehead ; side whiskers on face 
light huff to cream-huff. (Northwestern Costa Rica and higlier 

p<irtioiis of northern Nicaragua.) frontatus 

JJ. Chest and belly normally neither tawny nor orhrnceous-rufous. 

J/. Upper back distinctly darker than lumbar region of back; top of 
head. nJTk and upper back, and outer surfaces of fore and liind 
llmb.s varying from blackish to brownish black. 

002420—44 -2 



10 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

N. Underi>arts normally silvery whitish ; brownish black of head and 
neck passing gradually through cinnamon-drab on upper back 
to near olive-buff on lumbar region, the light drab extending 
along median line to base of tail ; side whiskers on face whit- 
ish. (Quintana Roo, Yucatan, and northeastern Guate- 
mala.) yucatanensis 

NN. Underparts not normally silvery whitish. 

O. Upperparts dark, especially on head, neck, and upper back, and 
more uniformly overlaid with black ; lumbar region more or 
less noticeably suffused with cinnamon ; pelage typically 
longer and denser ; underside of neck and throat mixed brown- 
ish and buffy ; rest of underparts including at least inner sur- 
faces of upper portions of limbs pinkish buff to cinnamon-buff, 
becoming cinnamon or rusty along sides of body ; crown cap 
blackish ; side whiskers on face blackish. (Alta Vera Paz, 

Guatemala.) pan 

00. Upperparts lighter, distinctly bicolored ; head, neck, and 
upper back brownish black ; lumbar region cinnamon-buff 
or cinnamon, darkened along midline by overlying blackish 
hairs ; underside of neck and throat varying from grayish 
to a mixture of buff-gray and brown ; underparts, includ- 
ing inner surfaces of arms in a strip narrowing from 
armpit to a point near elbows and on inner surfaces of 
legs to near ankles, varying from pinkish buff to cin- 
namon, becoming near cinnamon on sides of body ; crown 
cap blackish ; side whiskers on face dull whitish or yellow- 
ish. (Honduras and El Salvador northward through 
Guatemala to southeastern Oaxaca and the Isthmus of 
Tehuantepec and along east coast of Mexico to eastern 

San Luis Potosl.) vellerosus 

MM. Upper back not distinctly darker than lumbar region of back ; 
general coloration of upperparts, including outer surfaces of 
fore and hind limbs, as well as upper surface of tail, dusky, 
the darker hairs nearly sooty black on distal two-thirds and 
old gold on basal third, and thinly interspersed with these 
on head, neck, back, thighs, and upper surface of tail are 
old gold or silvery hairs ; coloration of upperparts further 
modified by lighter hair bases showing through ; a partially 
concealed huffy spot on forehead ; hairs on belly, inner sur- 
faces of hind limbs, and under surface of tail dull cinnamon- 
buff tipped with sooty black. (Rio Tuyra Valley of eastern 
Panama.) grisescens 



THE SPIDER MONKEYS KELLOtIG AND tlOLDMAX 11 

ATELES PANISCUS PANISCUS (Linnacug) 

Guiana Black Spider Monkey; Le Coaita 

[Siniin] pdiii.srus L^^•NA^n^s, Systoiiia natural, cd. 10, vol. 1, p. 2(5. 1758. 

SdiJitjoH pani'^cufi L.vcia'KDt; Tableau dcs divisicms, sous-divisions, ordres et genres 
lies inaiimiifOres. p. 4, linit. [Published as supplement to Di.scours d'ouver- 
turo et de cloture du cours d'histoire naturelle, et tableaux nu'thodique des 
inamniifdres et des oiseaux. ] 

Ateles pentadactyltis E. Gewfbroy, Ann. Mus. Hist. Nat. Paris, vol. 7, p. 2G9, 180G. 
Type locality, Cayenne, P^rcnch Guiana. (Description based on female col- 
lected by Martin.) 

Atclea nubpcnfadarti/Iuii Desmakest, Encyclopedie methodique (Zoologie), Mam- 
nialogie, pt. 1, p. 73. 1^20. Typo locality, '"La Cuyane fran(;ai.se, la cote de 
liancet au P(>rou" (part). 

A[t€lc8] utrr F. Cuvieb, Ilistoire naturelle des mammif^^res, vol. 3. livr. X). p. [1 I, 
pi. — , Mar. 1823. Type k>cality, Cayenne, French Guiana (see I. Geofifroy, 
1851, Catalogue methodique de la collection des mammif«^res, pt. 1 (Catalogue 
des Primates), p. 48). ("Le Cayou" is based on a rather young female.) 

[Cebus paniscus] siirituiDirnsis Fiscin^. Synopsis niammalium, p. 39, 182f). Type 
locality, not designated. 

[Cebus panismi] ftnioincnsis Fischer, Synopsis mammaliuni, p. 30, 1820. Type 
locality, "Guyana Gallica." 

Ti/pe locality. — "South America : Brazil" ["La Guyane" given as the 
"Patrie" by E. Geoflfroy, Catalo^ruo Mammi feres Mus. Nat. Hist. Nat. 
Paris, p. G, 18U3. Here restricted to French Guiana.] 

Type specimen. — Unknown. 

Dt.strihution. — North side of Amazon River east of Manaos and 
north to Caribbean coast of the Guianas. 

General characters. — A black monkey Avilli face normally flesh- 
cr)hjred ; pehi^e silky, lax, and very long (majority of hairs on mid- 
line of back 75-150 mm. in length, and on upper basal half of tail 
50-100 mm. in lenglh) ; hind foot large (190-220 mm.); tail often 
more than t\vice length of head and body. Differs from chamek of 
Peru mainly in normally flesh-colored instead of normally black face 
and in longer pelage (majority of hairs on midline of back less than 
75 mm. in length and on iipjwr base of tail less than 50 mm. in length 
in cham/'h). Differs from A. hehehuth hclzelntth of Venezuela in 
much longer pelage and nearly uniform black instead of contrasting 
colors of upi)er and un(lerf)arts. Differs from A. hehehuth maryinafus 
')f the Kio Tapajoz region of the southern side of the Kio Amazonas, 
Hrazil, in longer pelage, larger foot, and lack of the semilunar white 
forehead patch present in maryinatim. 

( ''ilor.--VjU\ iic pelage deep ghtssy black ; face noiinall y (lesh-coloied. 

t</.ull. — ^kull large, with anteiior profile strongly inclined backward 
to vault of brain ca.se; premaxillae, when viewed from above, rela- 
tively brf)ad ; nasal opening much elongated. Very similar to skulls 
of chamek and helzebrith^ but brain case somewhat less highly arched 
than either; rostrum bioailer. the sides more inflalid at base and 



12 



PROCEEDINGS OF THE NATIONAL MUSEUM 




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THE SPIDER MONKEYS KELU)GG AND GOLDMAN 13 



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PROCEEDINGS OF THE NATIONAL MUSEUM 



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THE SPIDER MONKEYS KELLOGG AND GOLDMAN 15 

sloping' uiilward more ^n-;ulually to zy<i()inata; maxillao fonninj; a 
more distinct ridpe behind posterior molars; thin oxtoinal redupli- 
cations of pteryijfoids cxtciidiii^ farther i^osteriorly. the posterior bor- 
ders less deeply concave behind tiie internal spinelike hanuilar proc- 
esses; posterior plane of posterior upper molars barely reaching ante- 
rior plane of temporal fossae. 

Mcasure77wnt,'<. — Two adult females from Rio Janiunda, near Faro, 
north bank of Rio Amazonas, Para, Brazil, respectively : Total length, 
i;330. 1.389 nmi. ; tail, 8T0. 920; hind foot, 195, 195. Two adult females 
from Lago Cuipeua, north bank of Rio Amazonas, Para, Brazil, respec- 
tively: Total length, 1,450, 1,413; tail, 880, 753; hind foot, 220, 220. 
Skull: Two adult females from Rio Jamundti, near Faro, Brazil, respec- 
tively: Greatest length, 118.8, 11G.8; orbital width, G8.3, G3.C; post- 
orbital constriction, 48.8, 51.3; width of brain case, 64, 63.3; zygomatic 
width. C8.7. 66.7; maxillary tooth row, 29.9, 29.5. Two adult females 
from Lago Cuipeua, Brazil, respectively: Greatest length, 113.6, 122.6; 
(jrbital width, 60.2, 68.2; postorbital constriction, 51.4, 51.4; width of 
brain case, 64.5, 64.6; zygomatic width, 69.4, 70.5; maxillary tooth row, 
29.9, 29.5. 

Remorks. — Linnaeus, in tlie tenth edition of Systema Naturae (vol. 
1, J). 26, 1758) combined the previously published descriptions of a 
spider monkey and of a howler monkey in formulating his diagnosis of 
ySimia^ paniscus. The spider monkey is described in the following 
words: "Simla fusca major, palmis tetradactylis, cauda prehensili ad 
apicem subtus nuda," and this is a word-for-word quotation from the 
account of the "four-fingered monkey" published by Browne (The civil 
and natural history of Jamaica, p. 489, 1789; edition of 1756 not con- 
sulted). The longer diagnosis at the end of Linnaeus's account of 
prini.scus seems to be based mainly on the accounts of the "Guariba" 
publi.shed by Marcgrave (Historiae rerum naturalium, p. 226, 1648; 
and Ilistoria natuial do Brasil, p. 226, 1942) and by Ray (Synopsis 
melhodica animalium quadrupi'«hnn et serpentini generis, p. 153, 
1693), .supplemented by additional anatomical details which are 
credited to Hallman and Aymen. 

In thf twelfth edition, however, Linnaeus (Systema naturae, ed. 12, 
vf)l. 1. J). 37, 1766) seems to have recognized that (he above-mentioned 
combination was inaccurate, and of the tenth edition references he 
cited only that of Browne (op. rit., 1756, |). 189) in the synonymy of 
[Shnia] paiihcuJi. With the exception of "Pedes Sc Cau(hi dimidia 
exterior bnninea,'' this revised diagnosis ai)|)lied to the "coaita." Fur- 
thermore, in the twelfth edition Linnaeus {i>p. r/'f., p. 37, 1766) i)laced 
the references to Marcgrave and Kay. which had l)een cited under 
paniy(ru.t in the tenth edition, in the syii'inymy of the howler monkey, 
[,S/////V/. I htlzehul. We are therefore of the o[)inion that Limiaeus in 



16 PROCEEDINGS OF THE NATIONAL MUSEUM vol.98 

1766 should be regarded as the first reviser and that the references to 
Marcgrave and Ray should definitely be eliminated in reaching a con- 
clusion concerning the status of paniscus. 

The "Guariba" of both Marcgrave and Ray is unquestionably the 
howler monkey, as was recognized by both Pennant (Synopsis of 
quadrupeds, p. 122, 1771; "preacher") and Goldfuss (Schreber, Die 
Sfiugthiere, Theil 1, Heft 7, p. 112, 1774) . The bases and type localities 
of the mammalian species in the tenth edition of Linnaeus were con- 
strued as they were by Thomas (Proc. Zool. Soc. London for 1911, 
pt. 1, p. 127, Mar. 22, 1911) on the assumption that Linnaeus's "quota- 
tion of his own earlier writings should be given absolutely overriding 
importance" and that all others should be ignored. The Linnean 
quotation under [^Simia] j^cmiscus in the tenth edition is "S. caudata 
barbata, cauda prehensili, palmis subtetradactylis. Syst. nat. 3," and 
on referring to the sixth edition of Systema Naturae (p. 3, 1748) , it will 
be noted that Linnaeus altered his original concept of this species by 
adding "palmis subtetradactylis," since the diagnosis under "/S'^m^(^" 
in the earlier edition reads "14. Simia caudata barbata, cauda pre- 
hensili. Marcgr. bras. 226." On the assumption that the reference to 
Marcgrave was the primary reference, Thomas designated Pernambuco 
as the type locality for paiiiscus. In view of the fact that Marcgrave's 
account of the "Guariba" applies solely to the howler monkey, the 
designation of Pernambuco as the type locality for paniscus cannot 
be accepted. Browne {op. cit., p. 489, 1789) stated that the "four- 
fingered monkey" is an inhabitant of the main continent, Linnaeus in 
the tenth edition gave the habitat of paniscus as "America meridionali : 
Brasilia," and in the twelfth edition after the removal of the references 
to the "Guariba" Linnaeus restricted the habitat to "America meri- 
dionali." Since "La Guyane" has been designated by Geoffroy {op. 
cit., p. 6, 1803) as the "patrie" of paniscus, we hereby restrict the type 
locality to French Guiana. 

The Guiana black spider monkey is readily distinguished from all 
others by its long, lax, silky pelage and its large foot. It is believed 
to be restricted to the area between the north bank of the Amazon River 
and the coast of the Guianas. It was met with by various early trav- 
elers, and several names proposed seem clearly to belong in synonymy. 
In the three specimens from Rio Jamunda, on the north bank of the 
Amazon, near Faro, Brazil, vestigial thumbs are present as pointed 
out by Tate (Bull. Amer. Mus. Nat. Hist, vol. 76, p. 215, Oct. 20, 
1939). In two from Lago Cuipeiia, also on the north bank of the 
Amazon, farther to the east, vestigial thumbs are present in the 
skeleton, as usual in the group, but do not appear to have been dis- 
cernible in the skin. Tate {op. cit.) refers to "small, triangular 
pterygoids without pointed tips" as a cranial character in paniscus. 
The external reduplications of the pterygoids are broken off in the 



THE SPIDER MONKEYS — KELLOGG AND GOLDMAN 17 

specimens examined by Tute, but the thin pterygoid plates in these 
and in other specimens from that region are in reality larger and 
more extended posteriorly than usual in spider monkeys. Other char- 
acter ascribed by him to paniacus are the small postglenoid processes 
and unexpanded zygomata of nearly uniform depth. Examination 
of large series of spider monkey skulls shows that these cranial fea- 
tures are everywhere subject to about the same wide range of individual 
variation. 

In DiUch (luiana, Kappler (Popular Sci. Monthly, vol. 32, No. 3, 
pp. 397-31*8, Jan. 1888) states that this monkey does not occur on the 
coast and that it is found only in the higher lands. An adult male 
collected by Kappler along the Marowijne lliver [ = Rio Maroni], 
Dutch (iuiana, is listed by Jentink (Catalogue systematique des 
maiiiiniferes, Mus. Hist. Nat. Pays-Bas, vol. 11, p. 41, 1892). Thomas 
(Ann. Mag. Nat. Hist., ser. 8, vol. 6, p. 505, Nov. 1910), however, 
records this monkey from the River Supinaam, lower Essequibo, 
Demerara. British Guiana. Richard Schomburgk (in Roth, Richard 
Schomburgk's travels in British Guiana 1840-1844, vol. 2, p. 72, 1923), 
states that this monkey is found mostly in troups of 16 to 20 in the 
higlust trees and that one was killed in the vicinity of Maripa at the 
forks of the upper Rio Waku-wau, a tributary of the Rio Tacutii, 
south of the Kaniiku Mountains, British Guiana. 

In French Guiana, Menegaux (Bull. Mus. Hist. Nat. Paris, vol. 8, 
No. 5, p. 21JG, 1902) has recordetl this si)ider monkey from the Riviere 
Camopi, a tributary of the Rio Oyapock, and from the Rio Lunier, 
a tributary of the Rio Carsevenne [=Calsoene]. 

Lonnberg (Arkiv for Zool., vol. 32A, No. 25, p. 8, july 18, 1940) 
has recorded specimens from the Igarape [==Rio| Anibii, north bank 
of Rio Ama/.onas opj)()sile mouth of Rio Madeira, and from Patua 
(or Paitana;, Pani, Brazil. 

apecimen^ examined. — Total number, 5, as follows: Buazil: Lago 
Cuipei'ia, noi-th bank Riu Amazonas, I'aia, 2 (M. C. Z.) ; Rio jMniuii(l;i 
near Faio. iioi I h bank of Ri(» Amazonas, Para, 3 (A. M. \. H.). 

ATKLKS PANISCL'S ( HAMKK (Humboldt) 
Bl^CK-FACED Bl^CK Sl'IDKR MO.NKKV 

isimiii chnnii.h lli MiiM DT, Hfciu-il (r<il)s«'rv!il ions tie '/uolnf^ic ct d'aiintoiiiic 

cuiiipur^f, vol. 1. J). .'{.'),'{, ISlli. 
AtclcH UiiiiiivuinhriH .J. A. Ai i KN. F?iill. Aiiht. Miih. Nut. Hist , vnl. WW, \\. dril, 

Yh'i'. 14, 1!M4. Type Inrjilily. Hiirirn. dc M.lKin.o, licMlwattTs of Uio Cy- 

raraii.'i, .Matto (Irosso, I?ruzil. 
[AtelcH atrr\ pcruvianux Lii.NNBFJW, Arkiv Mr Zool.. vol. .'{2A. No. 2.">, p. 1."., .Inly 

18, 1U40. TjjM? locality, ca.stfrti Peru. 

Ti/pe localitij. — Peru. [Here re.stiicitMJ to Uio Coiiilirniato, a 
tributary of Rfo Urubamba. Cnzco. Pciii.l 

002420 44 3 



18 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

Type specimen. — Unknown. 

Distribution. — Western Matto Grosso, eastern Bolivia, and north- 
eastern Peru to the Rio Solimoes and Rio Jurua, Amazonas, Brazil. 

General characters. — Entirely black, with pelage of moderate 
length, the hairs on midline of back 75 mm. or less, and on upperside 
of tail 40-65 mm. Similar in general to paniscus of French Guiana, 
but face black instead of flesh color; pelage shorter, the majority of 
hairs on midline of back 75 nxm. or less, instead of 75 to 150 ; tail often 
more than twice length of head and body and hind foot large as in 
paniscus; skull differs from that of paniscus in detail. Similar in 
color to A. fusciceps robustus of southeastern Colombia, but pelage 
entirely black without the short, sparse white hairs on chin and about 
the mouth of the latter that while inconspicuous appear to be impor- 
tant as a distinguishing character ; cranial details also different. Dif- 
fers from A. belzebuth beJzebuth of Venezuela in uniform black in- 
stead of contrasting black and buffy upper and underparts. 

Color. — Epidermis of face normally black and entire pelage deep, 
glossy black. 

Skull. — Similar in size to that of paniscus^ but brain case slightly 
more highly arched ; rostrum narrower, less inflated at base and slop- 
ing outward more abruptly to zygomata ; premaxillae more distinctly 
pinched-in behind incisors, the sides being almost parallel; maxillae 
more truncate instead of prolonged in a distinct ridge behind posterior 
molars; external reduplications of pterygoids less extended poste- 
riorly, the posterior borders more deeply concave; palate narrower; 
posterior plane of last molars reaching farther posteriorly into ante- 
rior plane of temporal fossae. Differs from that of robustus as fol- 
low^s : Premaxillae more extended anteriorly beyond canines, the sides 
more constricted and more nearly parallel at diastema between in- 
cisors and canines; nasal opening more elongated. Compared with 
that of belzebuth the skull is very similar in general, but the premaxil- 
lae differ in narrowness in about the same way as from robustus. 

Measurements. — An adult male and female from Rio Comberciato, 
Peru, respectively : Total length, 1,270, 1,380 mm.; tail, 820, 880; hind 
foot, 190, 202. Two adult females from Rio Yapacani, Bolivia, respec- 
tively : Total length, 1,320, 1,320 ; tail, 800, 800 ; hind foot, 220, 220. 
Skull: Adult male and female from Rio Comberciato, respectively: 
Greatest length, 117.5, 117; orbital width, 48.9, 50.8; width of brain 
case, 64.7, 59.8 ; zygomatic breadth, 68, 67.8 ; maxillary tooth row, 33, 
29. Two adult females from Rio Yapacani, respectively : Greatest 
length, 121.1, 118.8; orbital width, 63.8, 59.2; postorbital constriction, 
50.1, 48.7; width of brain case, 65.4, 64.7; zygomatic breadth, 71.3, 
68.2; maxillary tooth row, 31.4, 31.5. 

Remarks. — The large black spider monkey here treated as A. panis- 
cus chamek appears to be distinguished as the most completely black 



THE SPIDER MONKEYS KELLOGG AM) (JOLDMAX 19 

form ill tlie group. It afrroes closely in larfijc size with paniscus but 
dilFers in having black instead of flesh-colored face, shorter pelage, and 
cranial details pointed out. The general resemblance of chamek to 
A. fu^ciceps rohuyfiis of Colombia is very close, but the short white 
hairs on the chin and about the mouth, together with the smaller foot 
of the latter, are distinctive. 

Specimens collected by Johann Xalterer in 1825 on the Rio Sarare 
while en route from Matto Grosso (Villa Bella de Santissima Trini- 
dade) to Sao Vincente and in 1829 on the Rio Madeira above the mouth 
of the Rio Abuna probably are referable to this race (Wagner, Abh. 
math.-phys. CI. bayer. Akad. Wiss. Miinchen, vol. 5, Abt. 2, p. 418, 
1848). Similarly the specimens described by Miranda Ribeiro (Com- 
mi'^sao de Linhas Telegraphicas Estrategicas de Matto Grosso ao 
Aniazonas, Annexo No. 5, Hist. Nat., Zool., Manmiiferos, p. 8, May 
1914) from Cabeceiras do Pirocoluina and Jani on Rio Jarua, a tribu- 
tar}' of the G)--Parana, are likewise referred to this race. This spider 
monkey is rejiorted to occur also in the Serra dos Parecis in central 
Matto Grosso. The specimens mentioned by Lonnberg (Arkiv for 
Zool., vol. 32 A, No. 25, p. 10. July 18, 1940) from Puerto Salinas and 
Desierto on the Rio Beni, Bolivia, may belong here since three speci- 
mens (C. M. Nos. 2772, 2774, 2775) taken by Jose Steinbach on the Rio 
Yapacani, a tributary of the Rio Mamore, seem referable to this form. 

Under the name of Atcles ater \^=paniscus']^ Thomas (Ann. Mag. 
Nat. Hi.st., ser. 10, vol. 2, p. 251, Sept. 1928) listed specimens from 
Cerro Azul, altitude 2,000 feet about 35 miles west of Rio Ucayali, and 
from Chicosa, upper Rio Ucayali, Peru, about 35 miles below junc- 
lioii of I'rugamba and Tambo Rivers, altitude 1,500 feet. A similar 
allocation has been made i)y Thomas (Proc. U. S. Nat. Mus., vol. 58, 
p. 2:^:0, Nov. 10, 1920) of specimens (U. S. N. M. Nos. 194337-38 and 
-40) from llio Comberciato, a tributaiy of Rio Urubamba, Cuzco, 
Peru, altitude 3.O00 feet, and from Paciiitea. 

AccfU'ding to liartlett (Proc. Zool. Soc. L<jndon for 1871, pt. 1, p. 
218, June 1871) the black-faced spider monkey inhabits the forests on 
the Rio Ucayali and tlie low districts of the valley of the Ama/onas 
ill I*eru. A specimen from Peruate on Rio Maranon is recorded by 
Cabrera (Trabajos Mus. Nat. Cienc. Nat. IMadrid, ser. zool., No. 31, 
j». 1<'>. Oct. 28, 1!»17). A female obtained by Oastelnau and Devilje on 
tiie banks of the Jii(; Javari near its mouth on the Rio Amazonas is 
listed by I. Geolfroy (Catalogue methodique de la collection des mam- 
miferes, pt. 1 (('atalogue (h's I'rimales), [). 18, IS51 ) . 

It is possible that this bhick-faccd spider monkey may range down 
the Rio Maranon and the Solimoes to the Rio .Jurua in I>ia/,il since 
I^iinnlxMg (Arkiv f<ir Zool., vol. 32A, No. 25, \\ ]0, July IS, |!M0) lihts 
as ater .specimens from Lago (Jrande west of mouth of Rio Jurua on 
Rio Solimoes, from Jaburu and Joao Pessoa on the lower Hio Jurua 



20 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

and from Santo Antonio on the Kio Eiru, a tributary of the upper Kio 
Jurua. The specimens described by Von Ihering (Rev. Mus. Paulista, 
vol. 6, p. 409, 1904) from below Sao Felipe on the Rio Jurua should 
belong to the same race. 

Although Wallace (Proc. Zool. Soc. London for 1852, pt. 20, p. 108, 
May 23, 1854) remarks that another spider monkey, "probably Ateles 
oier^ inhabits the West Brazil district on the river Purus," recent work 
in Brazil has not confirmed this statement. 

Specimens examined. — Total number, 9, as follows : Bolivia : Ibon 
[ = Ivon], Rio Beni, 1; Rio Yapacani, 4 (1 skull only; C. M.) ; Santa 
Helena (upstream from Rurrenagaque), Rio Beni, La Paz, 1. Peru: 
Rio Comberciato, a tributary of Rio Urubamba, Cuzco, 3. 

ATELES BELZEBUTH BELZEBUTH E. Geoffrey 

Marimonda Spider Monkey 

Ateles helseluth E. Geoffroy, Ann. Mus. Hist. Nat., Paris, voL 7, p. 272, pi. 16, 
1806. 

Le chuva de Bracamorros Humboldt, Recueil d'observatious de zoologie et 
d'anatomie comparee, vol. 1, p. 8, 1812; "Ateles nuirginatiis," p. 341 (Prov- 
ince de Ja4n de Bracamoros, on the banks of Rio Santiago and Rio Ama- 
zonas [=Rio Maranon] betvpeen the cataracts of Yariquisa and of Pato- 
rumi, Departamento de Amazonas, northern Peru; specimen brought by 
Indians from Tutumbero, opposite Pongo de Cacangares, seen in house of 
governor at Tomependa, on Rio Maranon near mouth of Rio Chinchipe.) 

A[teles^ marimonda Oken, Lehrbuch der Naturgeschichte, Theil 3, Zool., Abt. 2, 
p. 1201, 1816. Type locality, Orinoco. 

Ateles fuligiiiosus Kuhl, Beitriige znr Zoologie uud vergleichenden Anatomie, 
Abth. 1, p. 25, 1820. Type locality, unknown. [Based on specimen in 
Mus. Nat. Hist. Nat. Paris.] 

Cebus hrissonii Fischer, Synopsis mammalium, p. 40, 1829. Type locality, RIa 
Orinoco, Venezuela. 

A\_teles'\ variegaius Wagner, Schreber's Die Siiugthiere, Suppl., Abt. 1, p. 313, 
1840. Type locality, Cocuy [=Cucui or Cucuhy], Rio Negro, northern 
Amazonas, Brazil, at the Venezuelan boundary (^dc Wagner, 1847, Abh, 
math.-phys. CI. bayer. Akad. Wiss. Miinchen, vol. 5, Abth. 1, p. 421) = 
Serra de Cocoi, upper Rio Negro {pie Sclater, Ann. Mag. Nat. Hist., ser. 4, 
vol. 6, p. 472, Dec. 1870). (Preoccupied by Simia variegatus Kerr, 1792= 
Ateles variegatus (Kerr), unidentifiable.) 

Ateles bartlettii Gray. Ann. Mag. Nat. Hist., ser. 3, vol. 20, No. 118, p. 300, 
Oct. 1867. Type locality, Brazil, the upper part of the Amazons ["Eastern 
Perti, near Xeberos"=Jeberos or Jeveros, Loreto (see Gray, Proc. ZooL 
Soc. London for 1867, pt. 3, No. 63, p. 992, pi. 47, Apr. 1868) =Forests of 
northwestern Perfi at highest point of mountains on forest trail between 
Moyobamba and Chayavitos, three days' journey from the latter (see 
Bartlett, Proc. Zool. Soc. London for 1871, pt. 1, No. 14, p. 217, June 1871).] 

Ateles chuva Schlegel, Mus. Hist. Nat. Pays-Bas, vol. 7, livr. 12 (Monogr. 40: 
Simiae), p. 175, 1876. Type locality, not designated. [Based in part on. 
"Le chuva de Bracamorros" of Humboldt, 1812 ; Ateles variegatus of Wag- 
ner, 1840; Ateles bartlettii of Gray, 1868; and two mounted female speci- 
mens, one from Peru, in the Leiden Museum.] 



THE SPIDER MONKEYS — KELLOGG AND GOLDMAN 21 

Atelcs prnblema Schtjcoel MS., in Jontink, Catalogue systi'matique dos mam- 
miferes, Mus. Hist. Nat. Pays-Bas, Leiden, vol. 11, p. I-, ]8;»2 {nomcn vudiiin). 

Type loeaT/fij. — I'nknown. [Iloro restricted to E>ineralda. west of 
(he mouth of Kio Giinpo, on Rio Orinoco, and south of JNIount Duida, 
Venezuela (see Humboldt, Recueil d'observations de zoologie et 
d'anatomie comparee, vol. 1, livr. 7, p. 3'2G, 1812).] 

Type .ypec'unvn. — A .specimen exhibited at public fairs until alter 
its death, when it passed into the possession of Reaumer, constituted 
the basis of "Le Belzebut" of Brisson (Rcfjnum animale, p. 211, 1756) ; 
this .stulFed specimen, the description of Builon, and two livin<i: spider 
monkeys in the menagerie were combined as the basis for Geotfroy's 
name. 

Distribution. — Ranges in the vast lowlands from near the junction 
of the Rio Orinoco and Rio Caura in central Venezuela* south to the 
valley of the Rio Xegro, westward to Colombia east of the Cordillera 
Oriental (Mambita), Ecuador east of the crest of the Andes, and to 
northeasteni Peru (Sarayacu). 

General characters. — Distinguished by unusual color pattern, the 
black upperparts contrasting strongly with i)ale buify undorparts, 
there being a sharp line of demarcation along lower part of sides. 
This marked contrast in coloration extends to the outer and inner sur- 
faces of its liml)s, although these vary in detail, and a triangular fore- 
head patch varying from white to golden or brownish yellow is nor- 
mally but not invariably present. Foot large (190-200 mm.) ; tail 
variable in length but often almost twice the length of head and body. 
Dilfers conspicuously in color, but similarity in cranial details indi- 
cates close relationship to hyhridus of Colombia. Differs notably from 
A. panlsru.<i poftiscu.s of French Guiana in much slioi-tcr polago and in 
contrasting colors of upper parts and underparts. 

Color. — Face and eyebrows black, a triangular forehead patch vary- 
ing from white to golden, yellow, or brownish yellow (occasionally 
entirely ab.'^ent or more or less hidden by stiff uj)t iirned black hairs over 
eyes and opposing black frontal tuft from crown) ; crown cap and 
upperparts generally black to a sharp line of demarcation with under- 
j)arts (in some individuals the black hairs under strong light have a 
.seal-brown tinge) ; underparts, in general, including inner surfaces of 
fore anfl hind limi)s, and underside of tail varying from cinnamon-bufT, 
yellow, or dull cream-buff to pale olive-buff; side whiskers on face, 
when present, whitish or buffy (absent in some specimens) ; ou(<'i- 
surfaces of foi-elimbs to elltows black; outer surfaces of forearms to 
wri.sts generally bbicjcish, liut occasioiuilly similar to undeiparts and 
intermixed with long dusky overhairs. Outer surface,s of liind limbs 
usually blackisli to knees or 1)elow, but in some specimens (j)articularly 
in eastern part of range) the outer suifaces of jiind limbs, as well as 
thighs and region arf)und base of tail, are distinctly st law-colored in 



22 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 96 

ground color thinlj^ overlaid with long black hairs (not numerous 
enough to materially affect the general ground color) ; hairs on throat 
normally black, sometimes .grayish, more or less concealed by black 
hairs ; tail normally bicolored, the upper surface black and the under 
surface similar to that of underparts (in occasional specimens the 
upper surface is about the same as that of the under surface, but more 
heavily interspersed with long black overhairs). 

Skull. — About as in that of hyhridus. Very similar in general to 
that of fanhcus but brain case somewhat narrower ; posterior borders 
of external reduplications of pterygoids more deeply concave. 

Measurements. — ^An adult male and female from El Llagual, Caura 
District, Venezuela, respectively : Total length, 1,257, 1,317 mm. ; tail, 
823, 810; hind foot, — , 193. An adult male and female from Valle de 
los Monos, Mount Duida, Venezuela, respectively: Total length, 1,110, 
1,330; tail, 695, 840; hind foot, 185, 195. Skull: An adult male and 
female from El Llagual, Venezuela, respectively: Greatest length, 
109.5, 117.7; orbital width, 60.2, 61.8; postorbital constriction, 50, 51; 
width of brain case, 59.7, 59.8; zygomatic width, 68.3, 66.7; maxillary 
tooth row, 29.8, 28.8. An adult male and female from Valle de los 
Monos, Venezuela, respectively : Greatest length, 119.4, 113.2 ; orbital 
width, 61.6, 61; postorbital constriction, 52, 52.1; width of brain case, 
61, 63.2 ; zygomatic width, 71.9, 68.4 ; maxillary tooth row, 29.9, 30. 

Remarks. — This well-marked and widely dispersed spider monkey 
has evidently been described under various names which appear to be 
assignable to the synonymy of hehehuth. Humboldt (Recueil d'obser- 
vations de zoologie et d'anatomie comparee, vol. 1, livr. 7, p. 353, 1812) 
observed that it was found along the banks of the Rio Orinoco but 
always above the Eaudal de Atures and the Raudal de Maipures in 
Venezuela. Allen records specimens from La Union on the Rio Caura 
(Bull. Amer. Mus. Nat. Hist., vol. 20, No. 29, p. 344, Oct. 8, 1904) as 
well as from the Rio Mato, a tributary of the Rio Caura (Bull. Amer. 
Mus. Nat. Hist., vol. 28, No. 12, p. 148, May 27, 1910) and from El 
Llagual on the Rio Caura, Venezuela (Bull. Amer. Mus. Nat. Hist., 
vol. 30, No. 10, p. 272, Dec. 2, 1911). Johann Natterer in February 
1831 obtained a male, three females, and a young of this spider monkey 
at Serra de Cocoi on the upper Rio Negro, Brazil, one female of which 
was exchanged with the British Museum of Natural History (Sclater, 
Proc. Zool. Soc. London for 1871, pp. 39, 225, 1871). 

According to Bartlett (Proc. Zool. Soc. London for 1871, p. 217), 
this spider monkey is found in Peru in the interior forest on the moun- 
tain range between Lamas (north of Rio Mayo, a tributary of Rio Hual- 
laga) and Saravacu on Rio Ucayali; on the lower spurs of the moun- 
tains between Moyobamba and the Rio Huallaga ; at Cahuapanas on 
the headwaters of the Rio Cahaupanas, a tributary of the Rio Ma- 
raiion; near the native village of Chamicuros (Parinari District) on 



THE SPIDER MONKEYS — KELLOGG AXD GOLDMAN 23 

the Rio Cliamicuros [ = Rio Siiniiria], a tiibiitai\v of the Rio Marafion; 
and on the Rj'o Ti<rre, a northern trihutarv of the Rio Maranon west 
of Nauta. It has been reported also from Puerto Indiana on the Ri'o 
Marafion just west of mouth of the Rio Napo by Tate (Bull. Amer. 
Mus. Nat. Hist., vol. 7G, p. 21G, Oct. 20. 1939), and from Elvira on 
the Rio Maranon above the mouth of the Rio Chanibira by Bartlett 
(Proc. Zool. Sec. London for 1882, p. 373), 

Sl)eciniens are also listed by Gooldi and Hajjjmann (Bol. Mus. Goeldi, 
vol. 4, p. 42, 1904) from Iquitos, on the Rio Marafion above mouth of 
the Rio Napo, by Cabrera (Trabajos Mus. Nat. Cienc. Nat. Madrid, 
ser. zool.. No. 31, p. 4G, Oct. 28, 1917) from Tarapoto (Nueva Floren- 
cia) on the Rio Nape, and by Loiniberg (Arkiv for Zool., vol. 14, No. 
4, p. 6, June 7, 1921) from below Baeza on the Rio Quijos, a tributary 
of the Rio Coca in eastern Ecuador. Festa (Boll. Mus. Zool. ed Anat. 
Conip. Univ. Torino, vol. 18, No, 435, p. 4, Feb. 11, 1903) identifies as 
Ateles variegatus several specimens collected by him in Ecuador in 
tlie valley of the Rio Santia^jo and at San Jose on the Rio Snni and 
remarks that he had found this spider monkey to be abundant in the 
forests bordering the Paute, Zamora, and Santiago Rivers. 

A specimen of this monkey (U. S. N. M. No. 3332) collected by 
William E, Moore in 1857 somewhere along the Rio Napo in eastern 
Ecuador has yellowish underparts and a golden brow patch. 

Spcehnens examined. — Total number, 9, as follows: Colombia: 
Mambita (Llanos) on Rio Guavio, a tributary of Rio Meta, 1 
(M. C. Z.); Villavicencio, western Rio Meta, 1. Ecuador: Rio Napo, 
1; Jinia (southwest of Sigsig) on headwaters of Rio Pamar, a tribu- 
tary of Rio Paute, 1 (skull only, M. C. Z.). Peru : No definite locality, 
1 (U. M. M. Z.). Venezuela: El Lhigual [ = Yagual], 2 
(A. M, N. H.) ; Valle de los Monos, Mount Duida, 2 (A, M, N. H,), 

ATELES BELZEBUTH MARGINATUS E. GcofTroy 
WniTE-WniSKERED SPIDER MONKEY 

Ateles maryinatuH E. GeomtOY, Ann. Mus. Hist. Niif. Paris, vol. 13, p. 92, pi. 10, 
Mar. 1809. 

AtelcH front (din Uhnnetit, Proc. Znol. Hoc. London for 1830-31, pt. 1, No. 4, p. 38, 
Apr. C, 1H;{1. TyiK' locality, unknown. 

Atclcti (ilbifroHH IL Sciiinz, Systcniail.sches Verzcicbni.ss aller bis jctzt bckannleu 
Siiugethlen? od»?r Synopsis Mamnialinrn, vol. 1, p. OS, 1841. [This name ap- 
pears In the Hynonymy of Atelca margitiatus and apparently Is based on 
Coaltn A fnmt blanc, AIcIch maryinatux Kr. (Jnvier, Ilistuiro nalun'll»> dcs 
niainniifcMcs, vol. 7, llvr. 01*, piiRcs untintMbi'rcd, Apr. is;{(). ) 

Type locality. — "Los parties dii I5iesil. iiiro.sees par le Rio- 
Janeiro" [=-Para, Brazil, and the borders of the Rio Orinoco, Vene- 
zuela; see E. GeofTroy, Ann. Mus. Hist. Nat. Paris, vol. 10, p, 100, 
1812. Here restricted to Cametfi, Rio Tocantins, Pani, Brazil. | 

Type specimen. — Museum National d'Histoire Natiirelle, Paris. 



24 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 96 

Distribution. — South side of Eio Amazonas, between the Rio Tapa- 
joz and Rio Tocantins, state of Para, Brazil. 

General characters. — A deep, glossy-black spider monkey, with a 
white semilunar forehead patch and a few white side whiskers on 
face. Similar in general to helzehuth., which it resembles in the pos- 
session of a conspicuous forehead patch, but the pelage is longer (long- 
est hairs on back about 90 instead of 50 mm.) and it differs notably in 
the extension of black over entire body and limbs. Differs from panis- 
cus of the opposite side of the Rio Amazonas in smaller size, especially 
the smaller foot, shorter pelage, and in the conspicuous white forehead 
patch (absent in paniscus) ; cranial details also different. 

Color. — An adult male from Caxiricatuba, east bank of Rio Tapajoz, 
Para, Brazil : Face flesh-colored, at least around eyes and on nose ; 
side whiskers white mixed with black; a narrow strip across brow just 
above eyes thinly clothed with erect black hairs behind which is a 
white semilunar forehead patch overlapped behind by black hairs from 
crown ; rest of upper and underparts deep, glossy black. 

Skull. — Very similar to that of hehehuth but interorbital region 
more depressed. Compared with that of paniscus the interorbital 
region is more depressed and the anterior profile rises more steeply to 
vault of brain case; it also differs notably in the lesser posterior exten- 
sion of the external reduplications of the pterygoids and the smaller 
winglike tips. 

Measurements. — An adult male from Caxiricatuba, east bank of 
Rio Tapajoz, Para, Brazil: Total length, 1,250 mm.; tail, 750; hind 
foot, 108 (188 ?). Three females from Marai, right bank of Rio 
Tapajoz as stated by Lonnberg (Arkiv for ZooL, vol. 32A, No. 25, p. 7, 
July 18, 1940), respectively : Total length, 1,114, 1,190, 1,245; tail, 770, 
613, 740; hind foot, 190, 189, 190. Skull: Adult male from Caxirica- 
tuba, Rio Tapajoz, Para, Brazil : Greatest length, 114.1 ; orbital width, 
61.3; postorbital constriction, 49.5; width of brain case, 59.6; zygo- 
matic breadth, 69.7; maxillary tooth row, 31.9, 

Remarks. — A. h. marginatu^s is readily distinguished by the white 
semilunar forehead patch combined with black general coloration. It 
appears to be most closely allied to helzehuth but differs in color pat- 
tern and in longer pelage. In length of pelage it suggests paniscus, 
which inhabits the opposite bank of the Rio Amazonas, but is less 
extreme, as the longer hairs on the middle of the back in that form 
exceed 100 mm. in length. In cranial characters also it agrees more 
closely with helzehuth than paniscus. 

This black spider monkey with white semilunar patch on forehead 
and white side whiskers seems to be most abundant between the Rio 
Tapajoz and the Rio Xingii. Specimens in the Museu Goeldi at 
Belem were obtained at Altamira on the Rio Xingu and at Santarem at 
the mouth of the Rio Tapajoz. Lonnberg (Arkiv for ZooL, vol. 32A, 



THE SPIDER MONKEYS — KELLOGG AND GOLDMAN 25 

Xo. 2"), p. T. July 18, 1040) iwords this monkey from Marai on the 
right bunk of the Rio Tapajoz, and Bates (The naturalist on the 
river Amazona, p. 217, IST;")) found it at Aracu on the Rio Cupary, a 
tributary of the Rio Tapajoz. It was found by Sieber at Cametii on 
the left bank of Rio Tocantins. 

Specimens e.vamhird. — Tt)tal number. 3, as follows: Brazil: Caxi- 
ricatuba, east bank of Rio Tapajoz, a few miles below Tauary, Pani, 
1 (M. C. Z.) ; no definite locality, 2 (1, C. M.). 

ATELES BELZEBUTH HYBRIDUS I. Geoffrey 

Hybrid Spider Monkey 

Atclcfi hi/hridiis I. Gvxhfroy, Mem. Mus. Hist. Nat. I'aris. vol. 17, p. 168. 1829. 

Aides albifrons (Jeay. Catalogue of monkeys, lemurs and fruit-eating bats in the 
collection of the British Museum, p. 44, 1870. Type locality. South America. 

[Atclcn bclscbnth] bnunieus Gr-w, Catalogue of monkeys, lemurs and fruit- 
eating bats in the collection of the British Museum, p. 44, 1870. Type locality, 
"Brazil." 

Amcr-anihropoidrs Joysi Montandon, Comptes Rendus Acad. Sci., I'aris, vol. 1S8, 
No. 11, p. S17, Mar. 11, 1929. Type locality, left affluent of upper Kio Tarra, 
a tributary of tiie liio Catatumbo, which flows into Lake Maracaibo, Vene- 
zuela. 

Type locality. — Valley of Rio Mao;dalena, Colombia. Here re- 
.<5trictcd to La Gloria on the Rio Magdalena, southern ]\Iagdalena De- 
j)artment, Colombia. 

Type specimen. — Cotypes, 2 male?5 and 1 female, Museum National 
d' Histoire Naturelle, Paris; presented by M. Plee in 182C. 

Distribution. — Serrania de Valledupar (Las Marimondas) and 
southward along the Rio Cesar to the Rio Magdalena (Puerto 
Estrella), and south in the Cordillera Oriental at least to latitude 6° 
N. in Santander (Bolivar), Colombia. 

General cluiraeters. — Distinguished by wood-brown general colora- 
tion of upperparts and a while triangular forehead patch. Apparently 
clo-sely allied to helzebuth of Venezuela, but contrasting strongly in 
brownish in.stead of black upperjiart.s. Differs from its geographic 
neighbor, A. jiiaciceps rohuntuH., of western Colombia, in brownish up- 
perparts and white forehead patch instead of nearly all black colora- 
tion, larger foot, and cranial details. 

Color. — Face and upturned hail's on anterior part of forehead just 
above eyes black, partially concealing a white triangular forehead 
patch; coloi-ation of ui)perj)arts in general wood brown, darker on 
head and upper back, becoming lighter and near avellaneous on lumbar 
region and hips; outer surfaces of forelimbs, thighs, and upperside of 
tail wood brown; underparts, inner surfaces of fore and hind limbs, 
and under surface of tail whiti.sh or buify ; siilc whiskers on face vary- 
ing from whiti.sh or buffy to daik wood brown. 

Skull. — About as in hrlzrhuth. Similar to that of A. pani^sciis panis- 
cus but variable, the premaxillae in some individuals less ])roduced 



26 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

anteriorly and the nasal opening less elongated. Compared with that 
of A. fusciceps robustus the premaxillae tend to be more produced 
anteriorly beyond plane of canines and the nasal opening more elon- 
gated. 

Measurements. — Three adult males from K,io Guaimaral, Valledu- 
par, Magdalena, Colombia, respectively: Total length, 1,248, 1,238, 
1,211 mm. ; tail vertebrae, 750, 739, 741 ; hind foot, 177, 178, 172. Three 
adult females from same locality, respectively: Total length, 1,330, 
1,247, 1267; tail vertebrae, 856, 778, 809; hind foot, 185, 187, 187. 
Skull: Three adult males already mentioned, respectively: Greatest 
length, 113.9, 114.4, 116.6; orbital width, 60, 65, 63.3; postorbital con- 
striction, 50.9, 49.2, 49.4; width of brain case, 65.6, 59.7, 65.3; zygomatic 
breadth, 71.4, 728, 74.1 ; maxillary tooth row, 30.3, 30.9, 33.3. Three 
adult females already mentioned, respectively : Greatest length, 116.9, 
109.9, 110.7; orbital width, 64.8, 59.9, 61.3; postorbital constriction, 
50.3, 48.1, 49.2; width of brain case, 61.6, 60.7, 60.8; zygomatic breadth, 
70.2, 69, 69 ; maxillary tooth row, 32.2, 32.6, 29.2. 

Remarhs. — Ateles liybridus was based on specimens presented to the 
Paris Museum by M. Plee in 1826. They were collected in the valley of 
the Rio Magdalena, Colombia, but no definite locality was given. It 
is now known to be a well-marked form with an extensive range, and 
for precision we restrict the type locality to La Gloria, on the Rio 
Magdalena in the southern part of Magdalena Department, Colombia, 
where the animal appears to be typical. Ateles albifrons Gray was 
described from South America without definite locality. According to 
Elliot (A review of the Primates, vol. 2, p. 45, June 15, 1913) , the type 
in the British Museum is stated on the ticket to be from Medellin, 
Colombia. That important town was for many years a headquarters 
for dealers in natural-history specimens, many of which labeled as 
from there are known to have been collected elsewhere, and it is not 
improbable that the type of albifrons may have been taken at some 
other locality. At any rate, descriptions of the type agree so well with 
hyhridus that the name may be assumed to belong in synonymy under 
it. 

Another spider monkey, [^Ateles helsehuthl bininneus^ was described 
by Gray (Catalogue of the monkeys, lemurs and fruit-eating bats in 
the collection of the British Museum, p. 44, 1870) as the fourth color 
variety of helzehuth. The original description of this specimen, which 
was stated to have come from Brazil, is as follows : "Brown, or brown- 
washed grey; cheek, loins, and outside of the thighs whiter; chest, 
throat, inside of limbs pale grey ; crown, outside of limbs, and upper 
surface of tail darker brown." Philip Hershkovitz has recently exam- 
ined the type of hrunneiis at our request and writes that it is identical 
with hyhridus. The type has been exhibited as a mounted specimen 
and is considerably faded. 

In cranial characters hyhridus seems to be identical with helzehuth., 
and the similarity in color pattern strongly indicates that the two are 



THE SPIDER MONKEYS — KELLOGG AND GOLDMAX 27 

conspecific. On the other hand, the cranial details as ^vell as color 
ditFerences point to a departure from robu^tus, a j^eographic neighbor 
on the west. This monkey has been reported from as far south as 
Bolivar in the Cordillera Oriental of Santander Department. 

Montandon (La Nature, Taris, No. 2809, vol. 1, p. 440, May 15, 1929; 
Coiiiptes Rendus Acad. Sci. Paris, vol. 188, No. 11, p. 817, March 11, 
1929) has described a monkey under the name of Amer-anthvopoldes 
luysi, typifying an assumed new family Amer-anthropoidae, from a 
left aftlucnt of the u|)per Rio Tarra, a tributary of the Rio Catatumbo, 
which flows into Lake Maracaibo, Venezuela. The collector. Dr. Fran- 
cois de Loys, observed two individuals and killed the female, and al- 
though an effort was made to save the skin and the skull these were 
subsequently spoiled by humidity. The description was based on notes 
and a photograph (Montandon, Journ. Soc. Amer. Paris, new ser., 
vol. 21, pp. 18:3-19;"). pi. 5, 1929) of the animal. The absurdity of the 
conclusions reached by Montundon is pointed out in detail by Cabrera 
(Rev. Soc. Argentina"^Cienc. Nat., vol. 10, pp. 204-209, July 12, 1930). 
The animal photographed seems to be unquestionably an Afclcs with 
a trianguhir white patch on the forehead. Specimens examined by us 
from tlie San Calisto district of the upper Rio Tarra proved to be 
A teles hclzchuth hi/hndus, to which the name Amer-anthropoules loysi 
is here relegated in synonymy. 

Specimens examined. — Total number, 17, as follows : Colombia : La 
Gloria, Rio Magdalena, 2; Las Marimondas, eastern Andes, Fonseca, 
Magdalena, o: Puerto Estrella, Rio Magdalena, Magdalena, 2; Rio 
Guaimaral, Valledupar, Magdalena, 6; Rio Tarra, San Calisto, San- 
tander, 2. 

ATELES FUSCICEPS FUSCICEPS Gray 

Brown-headed Spider Monkey 

Atclfn fusrircps Gray, rn.c. Zool. Soc. London for 18(i5, pt. 3, No. 47, p. TA'.i, Apr. 

1806. 
Atcles fuscicepi Sclateu. Proc. Zool. Soc. London for 1872, pt. 2. No. 42, p. GGIi, 

pi. 54 (col.). Nov. 187"J. (Specimen collcftod |»y (Jlarcnce Buckley in Trans- 

undeun Ecuador.) 

Type locality. — South America. [Here restricted to Hacienda Chi- 
nipamba,near Pefiaherrcra (west of Ibarra), Tntag Disti-ict, Tnibabura 
Province, ncn-thwestern Ecuador; allilnde 1,500 meters.] 

Type specimen. — Briti.sh Museum (Natural History) No. . 

Di-strihufifni. — Pacific side of cordilleia of TOcuador. 

General characters. — A black or bnnvnish-black si)ider monkey, willi 
top of head more distinctly brownish than body; foot small (150-170 
mm.) ; pelage rather eoapse; tail variable in length, usually one-fifth 
or more longer than head and body, clothed witii hair of moderate 
length. Closely allied to rohimtus of Colomi)ia but more brownish, 
especially on the head. Compared with paniscus, especially as repre- 



28 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

sented by A. paniscus chamek of Peru, the brownish head is distinc- 
tive: other differences are the short whitish hairs on chin (chin nearly 
naked and white hairs absent in chamek) ; shorter, coarser pelage, the 
crown patch less projecting over forehead, the shorter-haired tail, 
smaller foot, and cranial details. 

Color. — Face and upturned hairs on forehead black; crown cap 
tawny-olive, yellowish wood brown, or with black hairs tipped with 
burnt umber; remainder of body, including limbs and tail, covered 
with black hairs tinged with burnt umber. 

ShuU. — Skull of medium size and proportions; premaxillae mod- 
erately extended anteriorly; rostrum rather short and nasal opening 
of medium length ; auditory bullae somewhat flattened. Skull similar 
to that of the related form rohustus. Compared with that of panisciis : 
Premaxillae less produced anteriorly beyond plane of canines and 
anterior nasal opening less elongated. 

Measure^nents. — An adult male and female from Hacienda Chini- 
pamba, near Penaherrera, Ecuador, respectively : Total length, 1,090, 
1,281 mm.; tail, 720, 770; hind foot, 160, li69. An adult female from 
Cordillera de Chilluri, Ecuador: Total length, 1,194; tail, 655; hind 
foot, 152. Skull: An adult male and female from Hacienda Chini- 
pamba, near Penaherrera, Ecuador, respectively : Greatest length, 
113.7, 117.2; orbital width, 63.6, 62.5; postorbital constriction, 50.4, 
48.9 ; width of brain case, 61.2, 61.7 ; zygomatic breadth, 73, 71.2 ; maxil- 
lary tooth row, 32.6, 32.2. An adult female from Cordillera de Chil- 
luri, Ecuador: Greatest length, 113.8; orbital width, 58; postorbital 
constriction, 47; width of brain case, 59.7; zygomatic breadth, 69.6; 
maxillary tooth row, 30.2. 

Remarks. — Ateles fusciceps was based on a specimen from an un- 
known locality. Another specimen of this well-marked monkey was 
collected by Clarence Buckley in Transandean Ecuador, as recorded 
by Sclater {op. cif.)^ who also published a colored plate. For pre- 
cision in regard to the type locality, however, the name is restricted by 
us to the form occurring at Hacienda Chinipamba, near Peiiaherrera 
(west of Ibarra), Intag District, Imbabura Province, northwestern 
Ecuador, where the altitude is given as 1,500 meters. 

Lonnberg (Arkiv for Zool., vol. 14, No. 4, p. 5, June 7, 1921) has 
recorded specimens from near Gualea (alt. 4,000 feet) and Santo 
Domingo de los Colorados (alt. 2,000 feet) on the upper tributaries of 
Rio Esmeraldas, northwestern Ecuador. 

Specim^n^ examined. — Totalnumber, 8, as follows: Echjador: Caro- 
lina Ibarra, Imbabura Province, 1 (skull only, U. M. M. Z.) ; Cordillera 
de Chilluri, 1 (U. M. M. Z.) ; Gualea (near), Pichincha Province, alti- 
tude 3,000-4,000 feet, 1 (M. C. Z.) ; Hacienda Chinipamba, near Pena- 
herrera, Intag District, Imbabura Province, 2 (U. M. M. Z.) ; Mindo 
(below), Pichincha Province, altitude 4,000 feet, 1 (M. C. Z.) ; Par- 



THE SPIDER MONKEYS KELLOGC! AND GOLDMAN 29 

amba, Imbabura Province, 1 (skull only, U. M. M. Z.) ; no definite 
locality, 1 (U.M.M. Z.). 

ATELES FUSCICEPS ROBUSTUS J. A. Allen 

Colombian Black Spider Monkey 

Atclcs robiintu.'! J. A. Ali.i;n, Bull. Aiiier. Mus. Nat. Hist., vol. 33, art. 43, p. 052, 

Dee. 14, 1914. 
Atclcs dwicnsiii Goi.uman, Proc. Binl. Soe. Washington, vol. 28, j). lUl, Apr. 13, 

1915. Type healliy, near head t>l' Kin Llnion, Mount Pure, eastern Panama ; 

altitude 5,200 leet. 

2ype locality. — Gallera, Department of Cauca, western Andes, Co- 
lombia; altitude, 5,0(J0 feet. 

Type specimen. — Male adult, skin and skull, American Museum of 
Natural History No. 32354, collected July 13, 1911, by Leo E. Miller. 

Diatiibution. — Western cordillera of Andes from southwestern Co- 
lombia northward on west side of Rio Cauca to eastern Panama 

(.Mount Pirre). 

General characters. — All black, except for a slight brownish tinge on 
forehead of one individual and a few inconspicuous white hairs on 
chin and about mouth; hairs on back harsh and of medium length 
(majority of hairs on midline 40-70 mm. in length) ; tail variable in 
length, occasionally nearly twice as long as head and body ; hind 
foot .-nuiil (100-170 mm.) ; tail hairs of moderate length (30-50 mm. 
on ui)perside) . CIo.sely allied to fiusciceps of Ecuador but more nearly 
uniform black in color, without the distinctly brownish head of 
fuHciceps. Similar in color to paniaciLs of northern Brazil, but with 
a few white hairs on chin (chin nearly naked and white hairs absent 
in paniJiCws) ; foot .^mailer; hair on back and tail shorter; skidl dif- 
fering in derail. DilFers from (/eoffroyi and sub.species of Middle 
America in nearly uniform black, instead of diverse coloration, vary- 
ing from light buff to fenuginous. 

< olor. — Face and entire pelage deep glossy black, exeei)t in sonn' 
.''I)eciniens having a slight browni.sh tinge on the forehead and a few 
^slliti.'<ll hairs on chin and about mouth. 

SL-uU. — About a.s in the related form fuJ^clceps. Vei-y similar to that 
nf paniscu-s. but pninaxijlao less produced anteriorly beyond plane of 
canines; anterior nasal opening less elongated. Closely resembling 
that of f/ro/frt^y/^ but rostrum longer; anterior jjiofile risinir h'ss 
steeply from ends of premaxillae; auditory hiillar u-iially more 
flattened. 

.)f'ffMi/r/ments. — Type ( f roni original description): Total length, 
l.-"JO mill.: tail. r;J0; hind foot, IGO. Two adult fenuile topotypes, 
respect iv«'ly: Total length, 1.150, l.LiOO: tail, GOO, 750; hind foot, 155, 
lOO. ^7•////,• 'l\vo adult female topotypes, respectively: Greate.st 
length. ll.-).3, 111.7; (»rhital width, 58.9, 62.2; postorbital constriction, 



30 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 98 

47.9, 44.7; width of brain case, 59.1, 57.2; zygomatic breadth, 65.9, 
69.1 ; maxillary tooth row, 28.8, 31.1. 

Remarks. — Ateles danensis was based on a single specimen from 
eastern Panama, exhibiting characters that prove to be inconstant 
in additional specimens examined, and the name must be relegated to 
the synonymy of rohustus. Despite the marked contrast in color be- 
tween this black form and the red monkey of eastern Panama, the 
agreement in nearly all cranial details suggests close relationship. 
The auditory bullae are usually more flattened in rohustus, but even 
this feature is not always distinctive. To the southward, on the other 
hand, close alliance with fmciceps is indicated by the slight brownish 
tinge on the forehead in at least one individual, and apparent identity 
in cranial details. Allen (Bull. Amer. Mus. Nat. Hist., vol. 35, p. 235, 
May 31, 1916) records a specimen of this spider monkey from Barba- 
coas, Narino Department,, southwestern Colombia. 

Gray (Ann. Mag. Nat. Hist., ser. 4, vol. 11, No. 66, p. 468, June 1873) 
has recorded '■'■ Ateles ater'''' from the vicinity of Concordia, Antioquia, 
Colombia. Elliot (A review of the Primates, vol. 2, p. 30, June 15, 
1913) under the same name lists Cerete on the Rio Sinii, Bolivar, 
Colombia, as a locality record. Festa (Boll. Mus. Zool. ed Anat. 
Comp. Univ. Torino, vol. 18, No. 435, p. 4, Feb. 11, 1903) likewise iden- 
tifies a female taken in forest near the Rio Lara, which empties into an 
arm of Golfo de San Miguel, Panama, as Ateles ater. Anthony (Bull. 
Amer. Mus. Nat. Hist., vol. 35, No. 20, p. 375, June 9, 1916) refers to 
two specimens collected at Tapalisa, Panama. 

It is therefore evident that this black spider monkey ranges from 
the Serrania del Darion (Cerro Pirre and Tapalisa) southward along 
the mountain ranges bordering the Rio Sinii in Bolivar to the moun- 
tain range west of the Rio Cauca (Concordia), and southward at least 
to southwestern Colombia (Barbacoas). 

Specimens examined. — Total number, 7, as follows: Colombia: 
Gallera (type locality), 2 (A. M. N. H.). Panama: Cituro, 1 (A. M. 
N. H.) ; Mount Pirre, 1; Tapalisa, 1 (A. M. N. H.) : Rio Bayano, 1 
(skull only, M. C. Z.) ; Rio Tuyra, 1 (skull only, M. C. Z.). 

ATELES GEOFFROYI GEOFFROYI Kuhl 

NiCARAGUAN SpIDER MONKEY 

Atele[s'\ geoffroiji Kuhl, Beitrage zui' Zoologie nnd vergleichenden Anatomie, 

Abth. 1, p. 2G, 1S20'. (Printed also on same page as "Ateles Geoffrey, viihi 

species inedita.") 
Ateles melanochir Dksmarest, Encyclopedie methodique (Zoologie), Mammal- 

ogie, pt. 1, p. 76, 1820. [Based on specimen in Mus. Nat. Hist. Nat. Paris.] 

Type locality, unknown. 
Ateles melanochir Sclater, Proc. Zool. Soc. London for 1875, pt. 3, No. 27, p. 419, 

pi. 48 (col.), Oct. 1875. 
Ateles melanocerciis Scht.egel MS., in Jentink, Catalogue systematique des niam- 

mif^res, Mus. Hist. Nat. Pays-Bas Leiden, vol. 11, p. 43, 1802 {nomen riudum). 



THE SPIDER MONKEYS KELLOGG AND GOLDMAN 31 

T'lpc local'itij. — Unknown [Here restricted to San Juan del Norte 
(Gi-eytown), Nicaragua; see '•'•AteJc^ hyhrulus''' Sclater, Proc. Zool. 
Soc. London for 1862, pt. 2, No. 12, p. 186, Sept. 1862 (specimen from 
Rio R.ina, Gorgon Bay. near San Juan del Xorte) : and Salvin In 
Alston, Biologia Centrali-Americana, Mammalia, vol. 1, pp. 9-10, Sept. 
1879.] 

Type specimen. — Female adult, jVIuscum National d'Histoire Natu- 
relle, Paris (menagerie specimen acquired in 1819; see I. (icollroy, 
Catalogue methodique de la collection des mammiferes, pt. 1 (Cata- 
logue des Primates), p. 49, 1851.) 

Distribution. — Coastal region bordering San Juan del Norte or Ma- 
rina Bay, southeastern Nicaragua; probably ranging across through 
the lowlands to the Pacific coast. 

General characters. — Distinguished by light bulY, overlaid with 
dusky tipped hairs, in general coloration; dark markings on head and 
limbs variable and inconspicuous as a rule. Most closely resembling 
frantatus of northwestern Costa Rica, but lighter and dark markings 
much more restricted. Contrasts strongly with omatus of eastern 
Costa Rica in light buff instead of rich rufescent coloration. Differs 
from .1. fusciceps rohiLstus of eastern Panama and western Colombia 
in diverse instead of nearly uniform black coloration. 

Color. — Face and eyebrows varjnng from blackish to a mixture of 
black and buffy hairs, those on the brows directed upward to form a 
thin ruff; top of head thinly overlaid with blackish or brownish hairs 
directed forward, the under color usually light buff. Crown cap 
gi'ading from black to buff tinged with brownish; stiff long black 
hairs above eyes concealing to variable extent the white or buff tri- 
angular forehead patch ; upperside of neck, entire dorsal area, upper- 
part of arms all around and of legs to ankles all around, except knees, 
light buff tliinly intermixed with long black or brownish hairs; nar- 
low areas usually extending 2 or 3 inches up and down over knees 
black, the hairs black to roots, but in one specimen the knee patches are 
limited to a few overlying dusky hairs; elbows and outer sides of 
forearms more or less distinctly blackish; throat and sides of face 
and neck silky light buff to light ochraceous-buff ; underparts dull 
liglil buff; hands and feet blackish; tail above about like back, below 
.somewhat paler along basal portion, but with a narrow line of dusky 
hairs bordering callosity jiear tip. In one specimen all the darker 
areas are reduced and the general pelage is suffused with liglit oclira- 
ceous-buff, iMH-oming near cinnamon-buff on inguinal icgioii. lu an- 
other individual small cinnamon-buir ai-cas appear on shouldei-s, inner 
sides o[ wrists, sides of feet, and n<'ar middle of underside oi tail. 

SkitJl. — Very similar to that of ornntxiH but apparently smalhr. 

Mra,'<vrrmrnt}<. — No external measurements available. Skvll : An 
adult and a subadult fenialo from Nicaragua, resi)ectively : Greatest 



32 PROCEEDINGS OF THE NATIONAL MUSEUM vol. »►: 

length, 103.7, 101.5 mm.; orbital width, 58.7, 52.7; postorbital constric- 
tion, 46.8, 45.9; width of brain case, 55.7, 56.2; zygomatic width, 61.6, 
56.8 ; maxillary tooth row, 28. 26.2. 

Remarks. — A. geoffroyi appears to be the first identifiable name 
applied to a spider monkey from Middle America. At the time it 
was described the locality from which the type specimen came was 
unlmown. Much general evidence, however, indicates that this is the 
so-called gray spider monkey, also described as A. melanoGhir, now 
known to inhabit southeastern Nicaragua, and that this general gray 
(really light buff) style of monkey may range across through the 
lowlands of the Rio San Juan to the vicinity of Lake Managua and 
Lake Nicaragua. It is interesting to note that a very young example 
of front atus as figured by Gray (Zool. Voy. Sulphur, vol. 1, No. 1, 
Mammalia, pt. 1, pi. 1, Apr. 1843) has essentially the same light-gray 
color pattern as that of the adult of geoffroyi. It is probable that 
the type of geoffroyi came from San Juan del Norte, and for preci- 
sion we restrict the name to the animal that occurs in the vicinity of 
that locality. A specimen from Rio Rana, Gorgon Bay, near San 
Juan del Norte (Sclater, op. cit., p. 186, 1862) was subsequently listed 
by Gray (Catalogue of monkeys, lemurs and fruit-eating bats in the 
collection of the British Museum, p. 43, 1870) as ''Ateles hyhridus" 
from "St. Juan, Nicaragua." A very good description of this spider 
monkey, based on field observations, was given by Salvin to Alston 
(Biologia Centrali- Americana, Mammalia, vol. 1, pp. 9-10, Sept. 
1879). Confusion of the Nicaraguan race with A. hyhridus by Gray 
was undoubtedly due to external resemblance, which is rather strik- 
ing; but cranial characters indicate close alliance with geographic 
neighbors that are markedly different in coloration. A specimen 
(M. C. Z. No. 29626) from El Valle del Anton, near the Pacific coast 
east of the Azuero Peninsula in Panama, agrees closely in general 
coloration with those from Nicaragua, but the black on the crown 
patch and on forearms is more extensive. Whether this animal was 
native to the locality or brought from some other region seems some- 
what uncertain. 

Speciinens examined. — Total number, 10, as follows: Nicaragua: 
Managua, 6 ; no definite locality, 4 ( 1, A. M. N. H.) . 

ATELES GEOFFROYI VELLEROSUS Gray 

Mexican Spider Monkey 

Ateles vellerosus Gray, Proc. Zool. Soc. London for 1865, pt. 3, No. 47, p. 773, 

Apr. 1866. 
Ateles veUerosus Sclateb, Proc. Zool. Soc. London for 1872, pt. 1, No. 1, p. 5, pi. 2 

(col.), June 1S72. (Believed by A. Boucard to have been procured near 

Acapulco, Mexico.) 
Ateles neglectus Reinhakdt, Vid. Medd. Nat. Foren. Kjobenbavn, ser. 3, vol. 4 

(1872), Nos. 6-9, p. 150, 1873. Type locality, Mirador, Veracruz, Mexico. 



THE SPIDER MONKEYS — KELLOGG AND GOLDMAN 33 

Atdcs tricolor IIoujstek, Proc. Bicil. Soc Washington, vol. 27, p. 141, July 10, 
1914. Type locality. Hacienda Santa Ktigonia. S miles north of Tapanatepec, 
southeastern Oaxacn. 

Type localify. — ?Biazil. [Here restricted to Mirador, about 15 miles 
northeast of Huatusco, Veracruz, Mexico; altitude 2,000 feet.] 

Type spccitncn. — No. . British Museum (Natural History). 

DistHbution. — Unbroken forests of Veracruz and eastern San Luis 
Potosi and southeastward! lhr()U<xh Tabasco, across the Isthmus of 
Teluiantepec in eastern Oaxaca, to Honduras and El Salvador, except 
for highlands of Guatemala. 

General characters. — A subspecies distinguished by combination of 
black or brownish-black top of head, neck, and shoulders, in contrast 
with buffy lumbar region, and pinkish-buff to cinnamon-butf under- 
parts. Differs from yucatafnensis of Quintana Roo in deeper buff 
underparts (underparts in yucatanensis silvery white or light buff). 
Differs from pan of Guatemala in greater contrast in colors of upper- 
parts (in pan the upper back and lumbar region are more nearly uni- 
form blackish in tone). 

Color. — Top of head, neck, shoulders, outer surfaces of fore and 
hind limbs, and forearms all around varjnng from black to brownish 
black, passing near middle of back into cinnamon-buff or cinnamon and 
extending across lumbar region and including hips, the latter colors 
somewhat darkened along median line by overlying brownish hairs; 
underparts in general, including inner sides of arms in a strip nar- 
rowing from armpit to a point near elbows and on inner sides of 
legs to near ankles, varying from pinkish buff to cinnamon-buff, be- 
coming near cinnamon on sides of body; underside of neck varying 
from grayish to a mixture of buffy-gray and brown; face blackish, 
but lips and small tufts near sides of mouth are didl whitish in some 
specimens; hands and feet black; tail above black or brownish black, 
mixed below with buffy Juiirs in a median lijio gradually narrowing 
from base to near callosity. 

SkxiU. — Rather narrow, but closely similar in general to those of 
the other subspecies. Comi)ared witli yurafanrvsf.s the frontal region 
of the skull is slightly flatter, less convex along median line anteriorly, 
as shown in profile. 

Mea.'<nrem/'nt!<. — Two adult males from Sania Kfigenia, Oaxaca, 
resi)ectively: Total length, 1,200, 1,260 mm.; tail vertebrae, 7G0, 705; 
hind foot, 17G, 178. Average of 7 adult females from Santa Kligenia, 
Oaxaca: Total length, 1,204 ( 1, 140-1, .'^01 ) ; tail vertebrae, 751 (700- 
7ft0) ; hind foot, 170 (17H-1H5). SJcnll (three adult males from Sanla 
Kfigenia. respectively): Greatest lengtli. IlO.l. 112.5, 110.9; orbital 
width, 60, 56.3, 57.2; post orbital const i-iet ion, 51.3, 47.6, 47; width 
of brain case, 59.3, .58.5, .59.6; /ygoniatic wi.Ith, 70.5, 63.9, 68.r); 
maxillary tooth row, 28.9, 29.9, 29.9. Average of 7 adult females 



34 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

from Santa Efigenia, Oaxaca: Greatest length, 108.2 (105.2-113.4); 
orbital width, 56.2 (54.5-58.8) ; postorbital constriction, 46.7 (45.5- 
48.7) ; width of brain case, 58 (55.9-60.2) ; zygomatic width, 63.5 
(59.2-66.9) ; maxillary tooth row, 28.5 (27.2-29). 

Remarks. — A. velUrosus was originally assigned by Gray to Brazil, 
but this appears to have been an error, as pointed out by Alston 
(Biologia Centrali-Americana, Mammalia, vol. 1, p. 10, Sept. 1879). 
By some subsequent authors, including Alston, the name was applied 
to spider monkeys from Mexico without definite locality. Inciden- 
tally, the plate illustration of Sclater {op. cit.) closely resembles this 
form. It represents a specimen believed by Boucard to have been pro- 
cured near Acapulco, Guerrero, where it could not have been native, 
as no monkeys occur in that general region. Under the name velJerosus 
Thomas (Proc. Zool. Soc. London for 1890, pt. 1, p. 72, June 1890) 
recorded specimens taken in Veracruz at the following localities : Raya 
de Boca Agustin, Misantla ; boundaries of Misantla and Jalapa ; and 
Hacienda de Tortugas, Jalapa. At the time A. tricolor from south- 
eastern Oaxaca was described (op. cit) by Hollister, veUerosus was not 
recognized by him, and the name A. neglect us of Veracruz was over- 
looked. The following year, however, Hollister (Proc. Biol. Soc. 
Washington, vol. 28, p. 142, 1915) regarded tricolor and neglectus as 
identical. The various descriptions of veUerosus apply well to the 
series of specimens now available from both Veracruz and Oaxaca, 
under which we, therefore, include both neglectus and tricolor as 
synonyms. For precision we would restrict the name veUerosus to the 
form known to occur near Mirador, Veracruz, the type locality, inci- 
dentally of the later name, neglectus. 

The range of subspecies veUerosus marks tlie northern limit of the 
group in Middle America. It inhabits unbroken forested regions 
from near sea level to about 4,000 feet altitude, including some of 
the areas of heaviest and most continuous rainfall in Mexico. William 
Lloyd, a field agent of the former Biological Survey, writing of the 
mammals of the vicinity of Victoria, Tamaulipas, in 1891, stated 
that "a monkey has been said to have been taken a little south of here, 
but the most authentic information says they are not found north of a 
point west of Escandon, rather more than 50 leagues south." It is 
possible that spider monkeys ranged north into southern Tamaulipas, 
but information obtained in 1898 indicated that the northern limit was 
then near Xilitla, southeastern San Luis Potosi. 

The German naturalist Ferdinand Deppe, in 1825, purchased a live 
Ateles in Alvarado, Veracruz, which had been caught by a Mexican 
about 20 hours distant from that city. Afterward, Deppe (Sclater, 
Nat. Hist. Rev., vol. 1, No. 4, pp. 508-509, Oct. 1861) while en route 
from Caxaia to Alvarado, observed a "great number" of Ateles in a 
forest near Valle Real. 



THE SPIDER MONKEYS KELLOGG AND GOLDMAN 35 

One specinien fn)iu Juchitan, Oaxaca, has silvery whitish under- 
parts, and another from the same locality is normally colored. A 
very y()un<2: individual from Santa Eli<r('nia, Oaxaca, does not differ 
aj>preciably in color from adults from the same locality. One very 
younjnr one from Tehuantepec has the upperparts normally colored, 
and another from the same locality is uiuisually daik. tlie luinl)ar re- 
gion nearly as dark as the upper back. 

Specimens examined. — Total numl)er, 59, as follows: Mexico: 
Oaxaca : Juchitsin, 3 (A.M.X.H.) ; Oaxaca, without delinite locality, 
4 (A. M. X. H.) ; Santa Eligenia, 13; Tehuantepec, 4 (one skull without 
skin. A.M.X.H.; one skull without skin, U.M.M.Z.) ; Tuxtepec, 2. 
Tahax'o: Teapa. 1. Veracruz : Barranca de Boca, Canton de Jalapa, 
1 (skin only) ; Cuatotolapan,2 (skulls only, U. M. M. Z.) ; Pasa Xueva, 
VJ (2 skulls without skins, A. M. N. H.). Hondukas: Tegucigalpa: 
Cantoral. 1 (A.M.X.H): Guaymaca, 2 (A.M.X.H.). Olancho: 
C'atacamas, 1 (A.M.X.H.). Ocotepeque : El Chorro, east side of 
Ocotepeque, 4.500 feet, 1 (skin only, A.M.X'^.H). El Salvador: San 
Mi^nicl : Lake Olomega [=Laguna Lomego], 5 (C. D.), 

ATELES GEOFFROYI YUCATANENSIS, new subspecies 

Yucatan Spider Monkey 

Tj/pe locaVti}/. — Puerto Morelos, northea.st coast of Quintana Roo, 
Mexico; altitude 100 feet. 

Type specimen. — Male adult, skin and skull; U. S. N. M. No. 108531 
(Tiiological Surveys collection) ; collecled April 2, 11)01, by E. AV. 
X(Is(jn and E. A. Goldman; original number 14052. 

Distribution. — Forests of the Yucatan Peninsula, northeastern 
Guatemala, and ])robably adjoining parts of British Honduras; 
doubtless intergrading to the .southward with vcUcrosus. 

General characters. — A rather small, slender, light-colored race, 
with entire underparts silvery whitish or very pale buff, pelage short 
and thin. Size about as in vrUiVoimfi of Veracruz but decidedly paler, 
especially on underparts where in typical specimens a whitish silvery 
tone extends to neck and inner sides of limbs; underside of tail 
'•ream-buff to near callosity; frontal outline of skull more i)rominent. 

Color. — Type: Top of head, neck, and outer surfaces of fordimbs 
brownish black, pa.s.sing gradually through cinnamon-drab and light 
drab on shoulders and anterior part of back to near olive-bull on 
hips, the light drab extending along nie<lian line to base of tail: fa<e 
mainly dusky, but cheeks whitish; knees brownish black: res( of outer 
surfaces of hind limbs cinnamon-drab; entire under|)arts inebHJing 
underside of neck, inner surfaces of limbs to wrists and ankles thinly 
haired, between crcam-buir and silvery whili.sh; hands and feet black- 
ish; tail cinnamon-diab above and on sides. becoming cream-bulf below 
along median line from base to near callosity. 



36 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

SktiJl. — Similar in size to that of veUerosus^ but frontal profile more 
convex anteriorly; dentition rather light. 

Measurements. — Type : Total length, 1,150 mm. ; tail vertebrae, 766 ; 
hind foot, 181. Two adult male topotypes, respectively : Total length, 
1,141, 1,090; tail vertebrae, 744, 817; hind foot, 167, 174. Three adult 
female topotypes: Total length, 1,17^, 1,150, 1,056; tail vertebrae, 756, 
766, 713; hind foot, 169, 171, 165. 

Skull (type and two adult male topotypes, respectively) : Greatest 
length, 109.7, 105.7, 107.8; orbital width, 57.2, 53.9, 55.6; postorbital 
constriction, 47.8, 44.8, 45.8; width of brain case, 59.5, 56.8, 57.3; 
zygomatic width, 65.3, 61, 66.3; maxillary tooth row, 27.3, 27.5, 28. 
Three adult female topotypes, respectively: Greatest length, 104.8, 
108.5, 104.8; orbital width, 52.8, 54.1, 53.1; postorbital constriction, 
44.6, 45.8, 45.6; width of brain case, 55.1, 57.2, 57; zygomatic width, 
58.3, 60.2, 60.3; maxillary tooth row, 25.7, 29.8, 28.7. 

Remarks. — The spider monkey of the Yucatan Peninsula is readily 
distinguished by the silvery whitish underparts. Specimens from 
Apazote, Campeche, and Uaxactum, Guatemala, are referable to the 
present form, but in slightly darker and more buffy underparts they 
indicate gradation toward veUerosiis. One topotype and a young 
without definite locality have sparse white hairs on the anterior part 
of the crown and a vestige of a white forehead patch partially con- 
cealed by stiff black hairs. 

Specimens examined. — Total number, 16, as follows : Mexico : Cam- 
peche : Apazote, 1. Quintana Roo : Puerto Morelos, 6. Guatemala : 
Uaxactum, 8 (U. M. M. Z.) . No definite locality, 1 (A. M. N. H.) . 

ATELES GEOFFROYI PAN Schlegel 

Guatemalan Spider Monkey 

Ateles pan Schlegel, Mus. Hist. Nat. Pnys-Bas Leiden, vol. 7, livr. 12 (Monogr. 
40, Simiae), p. 180, 1876. 

Type locality. — Coban, Alta Vera Paz, Guatemala. 

Type speehnen. — Cotypes, adult male and two adult females, 
Museum d'Histoire Naturelle des Pays-Bas, Leiden. 

Distribution, — Mountains of central Guatemala; doubtless inter- 
grading with vellerosus. 

General characters. — A very dark spider monkey with lumbar region 
only slightly, if at all, lighter than shoulders and thus lacking the 
contrasting dorsal areas presented by vellerosus and other Middle 
American races ; pelage long and dense. 

Color. — Face, top of head, shoulders, outer surfaces of limbs, fore- 
arms all around, feet, and entire tail black or brownish black. In one 
specimen the dark color extends nearly uniformly over all of the 
upperparts, but in two others the lower part of the back is suffused 
with cinnamon partially concealed by overlying dusky hairs; under- 



THE SProER MONKEYS — KELLOGG AND GOLDMAN 37 

side of neck and throat thinly covered with mixed brownish and buffy 
liairs; rest of underparts, incKiding a strip narrowing from armpit 
to a point near inner side of elbow, and inner sides of hind limbs to 
ankles between pinkish butf and cinnamon-bufT, becoming deep cinna- 
mon or rusty along sides of body. 

ShuII. — About as in vcllerosus. 

Mea-s'iircin-cnts. — No external measurements available. Skull: Two 
adult females from Guatemala, respectively: Greatest length, 1G9.1, 
lO.'i,!; orbital width, 57.3, 59.4; postorbital constriction, 48, 50; width 
of brain case, C0.*2, 62.4; zygomatic width, G6.8, 68.8; maxillary tooth 
row. 27.8. 27.5. 

Remai'ks. — Three specimens exhibited by the Guatemalan Govern- 
ment at the World's Columbian Exposition were presented to the U. S. 
National Museum. These Avere labeled ''Guatemala" without definite 
locality but are believed to be from Alta Vera Paz and probably repre- 
sent typical Ateles pan. Aside from the very dark coloration, the coat 
is long and heavy, indicating that they came from a high elevation in 
the mountains. The skulls indicate close agreement with vellerosus 
in size and general details. 

Specimens examined. — Total number, 3, as follow's: Guatemala: 
Alta Vera Paz, 3. 

ATELES GEOFFROYI FRONTATUS (Gray) 

Black-foreheaded Miriki or Spider Monkey 

EriodcH fronintun C;il.vy. Ann. Miig. N.it. Hist., ser. 1, vol. 10, No. G'), p. 250, 

Dec. 1842. 
lirarhj/tclrs frontatun Or,\y, Zoology of the voyage of H. M. S. Sulphur, vol. 1, 

No. 1, Maiumalla, pt. 1, p. 9, pi. 1 ( col. ) , Apr. 1843. 

Type locality. — South America [ = harbor of Culebra, Le6n = Cule- 
bra. Bay of Culebra, (iuanacaste, northwestern Costa Rica, fide Gray, 
Zool. Voy. II. M.S. Sulphur, vol. 1, No. 1, Mannnalia, pt. 1, p. 10, Apr. 
1843.] 

Type specimen. — Female adult wit h young shot by Capt. Sir Edward 
Hr-lcher; British Museum (Natural History) No. . 

Dint rihut ion. — Noi'tli western Costa Rica and cxtrcinc western and 
northern Nicaiagua. 

General cJuiracters. — Similar in j)att<'ni of coloration, that is, the 
normal restriction of bhuk areas to top of head and irregularly to 
•juter surfaces of limbs, to yeoffroyi of .southea.stern Nicaragua, but 
bo<ly darker, the ground coloi* of iipperparts near buckthorn brown 
or Mars l)rown and of underi)aits honey yellow to tawny, instead of 
light buir. DifTers frf)m p(rn<nnei)»iH of Panama in brownish instead 
of deep ferruginous general body color. Differs from vellerosv^ of 
Veracruz in the restriction of black areas on anterior ])ait of back and 
more yellowish tone of lumbar regic^n. 



38 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

Color. — Face, forehead, top of head, and in some specimens back 
of neck and shoulders black; anterior part of crown patch more or 
less suffused with cinnamon-buff, owing to light basal color of hairs, 
and tending to form a transverse band across forehead ; side whiskers 
on face light buff to cream-buff; upperparts in general buckthorn 
brown to Mars brown, sparsely or noticeably intermixed with blackish 
hairs (especially on midline of back in some individuals) ; belly honey 
yellow to tawny, extending downward on inner surfaces of legs to 
ankles, and a lighter tone near cinnamon-buff extending downward 
on inner surfaces of arms to wrists; outer surfaces of arms and legs 
usually black (in some individuals restricted more or less to elbow 
and knee patches) ; side whiskers on face light buff to cream buff; 
tail usually sharply bicolor, black or dusky above to tip and tawny 
below to callosity (when otherwise, mixed light and dark hairs cover 
the tail above and below) . 

Skull. — About as in fanmnensis., averaging somewhat broader than 
in geoffroyi or vellerosus. 

Meamremsnts. — No external measurements available. Skull: An 
adult male from Peiia Blanca : Greatest length, 109.3 ; orbital breadth, 
59.4; postorbital constriction, 49; width of brain case, 58.6; zygomatic 
breadth, 69.6; maxillary tooth row, 28.4. Two adult females from 
Lavala and Tuma, Nicaragua, respectively : Greatest length, 117.3, 112; 
orbital width, 64.2, 60.2; postorbital constriction, 51.1, 51; width of 
brain case, 62.8, 59.8 ; zygomatic breadth, 71.4, 65 ; maxillary tooth row, 
29.6, 28.4. 

Remarks. — A. geoffroyi front atus is assumed to be a recognizable 
subspecies, although aside from the type in the British Museum, from 
northwestern Costa Rica, little is definitely known of its characters or 
distribution. Certain specimens from Costa Rica without definite 
locality, and from various localities in Nicaragua, are tentatively re- 
ferred to frontatus., although in none of these is the black on the head 
so restricted as in the type as figured by Gray {op. cit.). The color 
pattern in this form suggests close relationship to geoffroyi^ which 
might also be expected on geographic grounds. A very young individ- 
ual (U. S. N. M. No. 61208) from Costa Rica, without definite locality, 
closely resembles Gray's figure just mentioned in light coloration. A 
skull of an old female from Lavala, Nicaragua (A. M. N. H. No. 
28419), is one of the largest examined from anywhere in Middle 
America. 

Specimens examined. — Total number, 38, as folloAvs : Costa Rica : 3 
(without definite locality) . Nicaragua : Lavala, 2 (skulls only, A. M. 
N. H.) ; Peiia Blanca, 3 (A. M. N. H.) ; Rio Siquia, 2 (skulls only, 
U. M. M. Z.) ; Rio Yoya, a tributary of Rio Princapolca, 25 (skulls 
only, A. H. S.) ; Tuma, 2 (skulls only, A. M. N. H.) ; Uluce, 1 
(A. M. N. H.). 



THE SPIDEK MONKEYS KELLOGG AND GOLDMAN 39 

ATELES GEOFFROYI ORNATUS Gray 

Costa Rica Ornate Spider Monkey 

Hrlcs ornatuK Guay, Cntalopiie of monkeys, It'inurs and frnit-catitif; l)ats in the 
collection of the British Museum, p. 44, 1870. 

7'i/pc locality. — Unknown. [Here restricted to Cual)ro, Tahunanca 
region, southeastern Costa Kica.] 

Type apecimen. — British Museum (Natural History) No. . 

Distribution. — Eastern sh)pe of central cordillera of Costa Rica; 
doubtless intergrading ^vith panamensis on Pacific side of the central 
mountain range. 

General characters. — A dark golden-yellowish subspecies, the upper- 
parts in strong light having a glossy, golden-yellow sheen, owing to the 
yellowish subterminal bands of hairs. Resembling panamensis of 
I*anama and western Costa Rica but brighter, more golden yellowish in 
tone, the back, however, more obscured by overlying black-tip])ed hairs, 
throat lighter, underside of tail and inner side of upper arm lacking the 
deep '"ferruginous" of panamensis. Differs from frontatus of Guana- 
caste, northwestern Costa Rica, in greater extension of black area on 
limbs and over back of neck and shoulders, and black-tipped hairs 
overlying and tending to modify the golden-yellowish tones on back. 
Contrasting strongly in golden yellowish, instead of light bully colora- 
tion, with geofroyi of the coast region of southeastern Nicaragua. 

Color. — Face, top of head, forearms all around, outer sides of legs, 
hands, and feet black; upperpai-ts from shoulders to hips and base of 
tail glossy golden-yellow or apricot buff, varying in some specimens 
to cinnamon-rufous, more or less overlaid with blackish hairs; chest 
near cinnamon-bull", the rest of underparts and Hanks becoming near 
tawny or cinnamon-rufous, this general color extending down along 
inner sides of legs in some sperimens nearly to ankles; throat and under- 
side of neck near pinkisli buff; tail black ai)<)ve, more or less mixed 
with tawny below from base to near callosity. 

S/ntll. — About like that of panamensis-^ similar to thai of geoffroyi., 
but apparently .somewhat larger. 

M eajturemenis. — No external measurement s ava i lable. Skull : Three 
adult males from Talamanca i-cgion. Costa Rica, resjx'ctivdy : Cxreatest 
length, 110.2, 111.2. 1 i:',.H mm. ; ori)ital width, r.:}.;^ (',1.7, C.'i; postorbital 
constriction, 45.2, 45.7, 49.0; width of brain case. Gl.O. 58. C1.7; zygo- 
matic bivadth, 71.5, CH.2. 71.0; maxillary tooth row, 27. G, 30.f), 27.9. 
Average of adult females from Talanianca region, Costa Rica: 
CM-eatest length, 108.1 (104.5-114.4); orbital width. C.l.G (5G.4-64.1); 
postorbital constriction. 47.4 (4-1.H-49.1 ) ; width of brain case, .5H.7 
(55.7-61.3); zygomatic width, GG.8 rG;'..7-70.7) : niaxilhirv tooth i-ow. 
27.5 (26.3-29..5). 

Rryturrkx. — Much confusion has existed regarding the status of this 
brightly colored sul)species. Specimens that we believe belong here 



40 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

have commonly been referred somewhat doubtfully to A. geofroyi 
Kuhl. The description of A. omatus by Gray {op. cit.), from an 
unknown locality, applies so well to specimens from eastern Costa Rica 
that their identity seems unmistakable. A more detailed description 
of the type than that published by Gray is given by Elliot (A review 
of the Primates, vol. 2, p. 45, June 1913) . The glossy quality as well as 
bright colors of the pelage suggests the propriet}^ of the name omatus. 
The contrast in color with plain buffy geo-ffroyl of southeastern Nica- 
ragua is very great, but close agreement in cranial characters and other 
evidence indicate probable intergradation. 

Specimens examined. — Total number, 14, all from Costa Eica as 
follows: Cataratas, Rio San Carlos, 2 (A. M. N. H.) ; Cuabre, Tala- 
manca region 1 (A. M. N. H.) ; Guapiles, Limon Province, 1 (A. M. 
N. H.) ; Santa Maria, 37 miles south of San Jose, 3 (skulls only, A. M. 
N. H.) ; Talamanca region, 7. 

ATELES GEOFFROYI PANAMENSIS, new subspecies 

Red Spider Monkey; Mono Colorado 

Type locality. — Cerro Brujo, about 15 miles southeast of Portobello, 
Province of Colon, Panama ; altitude 2,000 feet. 

Type specimen. — Female adult, skin and skull, U. S. N. M. No. 171489 
(Biological Surveys collection) ; collected June 8, 1911, by E. A. Gold- 
man ; original number, 21165. 

Distribution. — Forested regions of Panama east of the Canal Zone 
(Cordillera de San Bias) , and west through Chiriqui to central western 
Costa Rica. 

General characters. — A. rather large, deeply rufescent race. Very 
similar to omatus of the Caribbean slope of Costa Rica, but reddish 
tone more intense, the back less obscured by overlying dusky hairs; 
inner side of upper arm pinkish cinnamon to ferruginous. Differs 
from azuerensis of Azuero Peninsula, Panama, in deep reddish instead 
of cinnamon or tawny general coloration. 

Color. — Type: Upperparts from back of shoulders to base of tail, 
backs of thighs, and sides of body ferruginous, slightly obscured by 
black tips of hairs; outer surfaces of fore and hind limbs either black 
or blackish to knees and elbows ; outer surfaces of limbs below knees 
and elbows either black or with varying admixture of black and ferru- 
ginous hairs, the hairs banded but black at tips; underparts thinly 
haired, pinkish cinnamon on chest and along inner sides of hind limbs 
to ankles, becoming deep, clear ferruginous or dark dusty reddish over 
abdomen ; face, crown cap on head, and median streak on back of neck 
blackish ; sides of neck covered w^ith a mixture of pinkish buff, cinna- 
mon-buff, and blackish hairs ; chin nearly naked ; throat and under side 
of neck thinly clothed with light cinnamon-drab or light brownish 
hairs; tail above ferruginous heavily mixed with black on basal two- 



THE SPIDER MONKEYS — KELLOGG AND GOLD^LVX 41 

thirds, bec<)inin<r clear black toward tip, bolow deep ferruginous to near 
callosity where blackish hairs extend all around. The ji;round color 
of body in other specimens grades from dark rusty reddish to ferru- 
<:in(»us or burnt sienna, and a narrow ferrurrinous or cinnamon-buff 
streak usually extends long inner side of upper arm from armpit to 
elbow. 

/Sk-i/IL — Very similar to that of groffroyi, but brain case rather broad. 

Measurements. — Type: Total lenirth, 1.-2H0 mm.; tail vertebrae, 840; 
hind foot, 194. An adult male and ftinale from Cerro Azul, respec- 
tively: Total leuirth, 1.280. 1.225; tail vertebrae. 786, 785; hind foot, 
183, 176. Skull (type): Greatest length, 112.8; orbital width, 62; 
postorbital constriction, 52.4: wiilth of brain case, 61.2; zygomatic 
breatlth, 68: maxillary tooth row, '^7.3. An adult nuile and female 
from Cerro Azul. respectively: Greatest length, 114.2, 109.1; orbital 
width, 62.7, 59.4; postorbital constriction, 53, 50.3; width of brain case, 
63.2. 61.4: zygomatic width, 70.2, 69.6; maxillary tooth row 31, 28.1. 

/i'> nurrkft. — This is the monkey conunoidy known as ''mono colo- 
ra<l<i" in Panama and western Costa Rica. It is the most intensely 
reel of all the races of Middle America, but despite this it has been 
generally confused with the liglit bulFy form, geoffroiji, of Nicaragua. 
While the ground color of most of the body appears to be ferruginous 
to burnt sienna in ordinary light, in strong light and particularly in 
direct sunlight individual hairs on the upperjKirts are .seen to have 
glos.sy subtcrminal bands giving that part of the pelage a golden- 
yellowish sheen, a peculiarity still more pronounced in ornaPufi and 
not shared with the more northern races. The specimens from 
Chiriqui and as far west as central-western Costa Rica agi'ee closely 
in color with those from the (^anal Zone in the vicinity of the tyi>e 
locality. In one sjiecimen of a series of six skins from Pozo Azul, 
Costa Rica, a white patch on the forehead is nearly concealed in front 
by loiiL'er black-tipped hairs and the cheeks are dull whitish or ]>alo 
huffy. 

Specimens examined. — Totiil mimbci-, 21. as follows: Cosia Rir.v : 
Canas Gordas. Puntarenas, 3 (A. M. X. II.) ; Kl Pozo, 12 niiles above 
n)oiifb of Ri'o Grande de Terraba, 1 (slcin only, A. M. N. II.) ; Pozo 
A/nl. 10 miles above mouth of Rio Grande de Pirris, 12 (3, C. M.; 4 
skins with skulls and 5 extra .skulls, A. M. N. II.). P.\nam.\: Cerro 
Azid, near head of Chagres River, 2; Cerro Brujo {{\\)v locality), 1; 
Coto, Chiriqui, 2 (M. C. Z.) ; Rfo Pequeni, Canal Z(me, 2 (M. C. Z.) ; 
Snn Juan. Chagres River. 1. 

ATELE.S GROFPROVI A7.URRKNSIS Bole 

Azi'ERO Spider Monkey 

Atrlrs n:urrfn.«iif Hoik. S<1. I'lihl. flfvi-liiinl Mus. Nut. Illst., vol. 7, p. 149, AuR. 
.31. i;t.37. 



42 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

Type locality. — xVltos Negritos, 10 miles east of Monti jo Bay (part 
of the spur forming south drainage divide of Kio Negro), Mariato 
Suay Lands, Azuero Peninsula, Veraguas Province, Panama; alti- 
tude, 1.500 feet. 

Type specimen. — Female adult, Cleveland Museum Natural His- 
tory No. 1235. 

Distribution. — Forested mountains of the Azuero Peninsula, Pan- 
ama, probably in deeper forests on both slopes, but known only from 
the western (Veraguas) side from the vicinity of Ponuga southward. 
Possibly ranging west to Burica Peninsula on Panama-Costa Rican 
boundary. 

General characters. — A subspecies distinguished from neighboring 
forms by lighter tawny or ochraceous-tawny general coloration. 

Color. — Face blackish ; top of head and back of neck varying from 
deep black to a mixture of black and cinnamon-buff, with black pre- 
dominating; upperparts in general ochraceous-tawny thinly inter- 
mixed with black-tipped hairs; shoulders and outer surfaces of arms 
and legs black ; belly similar to back, but paler owing to absence of 
black-tipped hairs; throat and chest cinnamon-buff, this general tone 
extending downward to wrists and ankles on inner surfaces of limbs; 
tail above black, below tawny to proximal end of callosity. 

Skull. — Rather small and narrow, but very similar otherwise to 
that of panamensis. 

Measurements. — An adult male and female from type locality, re- 
spectively: Total length, l.lYO, 1,166 mm.; tail, 720, YOl; hind foot, 
177, 168. Skull (t3^pe, from original description) : Greatest length, 
110.8; width of brain case, 56.8; zygomatic breadth, 65.1; maxillary 
tooth row, 27.7. An adult male from type locality : Greatest length, 
109.8 ; orbital width, 57.7 ; postorbital constriction, 44.1 ; width of brain 
case, 58.7 ; zygomatic breadth, 67 ; maxillary tooth row, 29.7. 

Remarks. — This subspecies is definitely known only from the heavily 
forested mountains of the Azuero Peninsula, where it appears to be 
isolated, but may prove to have a more extended range near the Pacific 
coast of western Panama. A series of 25 skulls from the Burica Penin- 
sula, near the Panama-Costa Rican boundary, said to be from light- 
colored skins not saved and, therefore, not available for examination 
by us, are indistinguishable from those inhabiting regions as far north 
as Nicaragua. For reasons of geography and light color reported we 
refer them, tentatively, to as/uerensis. 

Specimens examAned. — Total number, 27, all from Panama, as fol- 
lows: Altos Negritos, Azuero Peninsula (type locality), 2 (CM. N. 
H.) ; Rio La Vaca, near Puerto Armuelles, Burica Peninsula, 25 (skulls 
only,A.H. S.). 



THE SPIDER MUX KEYS — KELLOGO AND GOLDMAN 43 

ATELES GEOFFUOYI GRISESCENS Gray 

Hooded Spider Monkey 

Atclfs griscscens Gray. rroe. Zool. Soc. Lnmloii for 1SG5, pt. 3, p. 732, Apr. 1806. 
Atil> s cumJlatu-^ Quay. Troc. Zool. Soe. London for l.S(>"), pt. 3, p. 733, Apr. ISfiO. 

Type locality, unknown. 
Atclvs oucullatiis Sclatix, rmc. Zool. Soc. London for 1871, pt. 1, p. 223, pi. 14 

(col.), June 1871. 
Alclct mclatiochir Sclater, Proc. Z<iol. Soc. London for 1875, pt. 3, p. 419, pi. 49 

(col.), Oct. 1875. 

Type 7o(?a7^7^/.— Unknown. [Here restricted to Rio Tuyra, south- 
eastern Panama.] 

7\)/pe si>eci77un. — Bi-iti.-«li Aruseum ( Xatural History) No. . 

D'.Mtributioj}. — Presumably the valley of Rio Tuyra and probably 
southeastward through the Serrania del Sapo of extreme southeastern 
Panama and the Cordillera do Baudo of northwestern Colombia. 

Gineral characters. — Adults apparently characterized l)y long, lax 
pelage and peculiar du.sky coloration, with a general admixture of 
yellowish gray or golden hairs, the hairs on upperparts golden at the 
base. The skull indicates close relationship to panarrieiisis^ despite the 
contrast in the deep reddish color of the latter. 

Color (type of nwuU-atufi). — Skin around orbits and on nose bare 
and of a browni.sh flesh color with darker freckles intermixed; cheeks 
and lower jaw nearly bare of hair, but .skin more decidedly of a black 
shade (Murie, Proc. Zool. Soc. London for 18G5, pt. 3, p. 739, Apr. 18G6) . 
Hairs very long and lax, those on head projecting forward over fore- 
head; crown and neck black; rest of body above and below, limbs, and 
tail black intermixed with yellowish-gray or golden hairs, so numer- 
ous as to give a pale yellowish-brown coloration to back (not far 
removed from color on the bases of the hairs in similar parts of A. 
f/riJiescens) ; hands and feet black, bases of hairs on hands golden, but 
not on those of feet which are black to the roots (Elliot. A review of 
the Primates, vol. 2, pp. 38-39, June 1913). Barbour (Journ. Mam- 
malogj'. vol. 13, Xo. 4, pp. 307-3G8, Nov. 1932) lias conunented on the 
.skin and skull of a spider monk(n' (M. C. Z. No. 27490) obtained at 
Chepigana in Darien. Panama. In this inunature specimen, the entire 
upperj)arts, including (he outer .surfaces of fore and hind limbs as well 
as the upl)er surface of the tail, are dusky, the darker hairs nearly sooty 
black on the distal two-thirds and thinly interspersed with these on 
head, neck, back, thighs, and upper surface of tail are old gold or silvery 
hairs; the cf)loration of the uj)perparts is further modified by a lighter 
sulTusion resulting from (he old-gold color of the basal third of these 
liairs showing through. A partially concealed bulfy spot is present on 
the forehead. Tlie thrf)at and chest are thiidy covered with sooty 
black hairs. On the belly, inner surfa<-es of hind limbs, and under 
surface of tail the hairs art- ilull ciniiamon-bufT tipped with sooty 



44 PROCEEDINGS OF THE NATIONAL MUSEUM vol.86 

black. The inner surfaces of fore limbs are noticeably lighter than 
the outer and also somewhat lighter than chest. The hairs on hands 
and feet are lighter at the base than at the tip. The hairs on the 
forehead project forward as on the type specimen. 

Skull. — Similar to that of panamensis. 

Measureme^its. — An adult male (type of cueuUatiis) : Head and 
body, 431±mm.; tail, 698; hind foot, 159 (fkle Murie, 1866, op. cit., p. 
T39). jSkuIl: Immature specimen from Chepigana: Greatest length, 
103; orbital breadth, 52.3; postorbital constriction, 47.6; width of 
brain case, 55.6 ; zygomatic breadth, 60 ; maxillary tooth row, 26.8.- 

Remarks. — A somewhat grizzled spider monkey obtained by E. 
Greey, an officer on the Eoyal West-Indian Mail Co.'s steamship 
Shannon at St. Thomas, Virgin Islands, for the Zoological Society of 
London became the basis of A. grisescens of Gray. After the death of 
the animal, the skin and skull were acquired by the British Museum 
(Natural History), and the skin was compared by Sclater (Proc. Zool. 
Soc. London for 1871, pt. 1, p. 223, June 1871) with the skin of a some- 
what similar half -grown male purchased from a London dealer in 1869. 
This specimen, also evidently of unknown native origin, he regarded 
as probably identical. The type of this spider monkey, according to 
Elliot (A Review of the Primates, vol. 2, p. 37, June 1913), is entirely 
black except for the yellowish-brown tint of the underside of the tail, 
but intermixed with the hairs of the upperparts are long, gray silvery 
or golden hairs. The latter are not sufficiently numerous to affect the 
general blackish coloration except on the shoulders, lower back, and 
the limbs where the hairs are yellow or golden at the base. 

The type of cucullatus^ a male now in the British Museum (Natural 
History), had a small tubercle representing the thumb, and Sclater 
{op. cit..f 1871, p. 224) remarked that he had "some reason to suppose 
it may be from the northern coast of Colombia, as ... a black spider 
monkey with long hair over its head is occasionally brought for sale 
into Cartagena." 

Attention is here directed to two black-handed spider monkeys fig- 
ured by Sclater {op. cH.^ 1875, pi. 49), which were received likewise 
from officers of the West Indian Mail Co. The color of the face and 
nose closely resembles that described for grisesoens^ and in addition the 
distal black color of the hairs on the upperparts does not entirely con- 
ceal the yellowish basal color of these hairs. If these specimens are 
correctly allocated to grisescens, it would seem that this spider monkey 
is subject to considerable variation in external appearance. Owing to 
the lack of adequate material for study the status of this form remains 
uncertain. 

Specimens examined. — Total number, 2, as follows : Panama : Che- 
pigana, Darien, 1 (M. C. Z.) No definite locality, 1 (yg., A. M. N. H.) . 



• Hindermost molar has not erupted. 



THE SPIDER MONKEYS — KELLOGG AND GOLDMAN 45 

INCERTAE SEDIS 

(?) ATELES RUFIVENTKIS Sclatcr 

Atih\s viUn-o.stci ScL-MKR, I'mc. ZuoL Soc. London for 1871, pt. 2, p. 478, Aug. 1.S71 

{tier Gray). 
Atrhs rufii'ftitrL'* Sci.ateu. Troc. Zool. See. London for 1872, pt. 2, p. G88, pi. 57 

(coL), Nov. 1872. 

Type locaJf'fr/. — Rio Atrato, Darien, Colombia. 

Type spi'chnrii. — Britisli Miisoinn (Xatural History) No. . 



Remarks. — Sclater (Ptoc. Z(K)1. Soc. London for 1871, pt. 2, p. 478, 
Au^. 1871) stated that the type of Afeles iniflventris^ a young female, 
with body length 12 inches, tail 15 inches, was obtained at Colon, Pan- 
ama, by an officer of the Royal West Indian Mail Co. and tJiat it had 
been brought from the Rio Atrato by one of the American party (pre- 
sumably the Dan'en E.xploring Expedition under the command of 
Comdr. T. O. Self ridge. U. S. X.) engaged on the survey of the isthmus 
for a ship canal. The color of this monkey was described as uniform 
black excejit for the flesh-colored face and muzzle and the pale fulvous 
belly; the color of the latter, however, hardly extending on the inner 
surfaces of the limbs. A colored illustration for this monkey was sub- 
sequently published by Sclater (Proc. Zool. Soc. London for 1872, })t. 
2, p. 688, pi. 57 (col.), Nov. 1872). Elliot (A review of the l^rinuit.es, 
vol. 2, p. 30, June 15, 1913), however, described this monkey as having 
the "underi)arts extending a short distance on inner side of arms and 
leg.s, bright rufous." and the line between the color of the underparts 
and the black of the body as sharply defined. 

Dt't<M-mination of the status of the name AfeJcs rufivnitrh presents 
some ditiiculty at the present time. The relative lengths of head and 
body to that of the .short-haired tail, the color pattern, and direction 
of hair on crown and foi-ehead corresp(jnd closely with these features 
in Ahtvattn paUiata ar<juatoriallH^ young females of which also lack 
tlio bushy beard present on the throat in the males. These resem- 
blances are shown very well by two .slightly larger young howlers 
{V. 8. N. M. Nos. 3349, 3359) also from the Rio Atrato. It .seems 
unlikely that representatives of both Ateles and Alouatta with the 
same color i)attern would be found in the same region. Sclater states 
that the thumb is absent, but examination of skins in the collection 
shows that the thumb in skins of some young individuals is incon- 
spicuous and might have been overlooked. The hand and feet in the 
illustration published by Sclater {ojk <it.) seem more clo.sely to re- 
semble those of AUmatta than those of Atrlrs. Elliot {op. c'li.) states 
that no skull .s^mmus to have been preserved. While all these points 
may be given consideration final generic allocation nuist await critical 
examination of the type, now not available. 



u. 1. «oviiii(MiNT fmNTiNa orrict: l>«4 



PROCEEDINGS OF THE UNITED STATE5 NATIONAL MUSEUM 



i33ueJ ff^jKyL Q*Ml hy the 




SMITHSONIAN INSTITUTION 
U. S. NATIONAL MUSEUM 



Vol. % Wathington: 1944 No. 3187 

A EEVISIOX OF THE AMERICAN CLIXGFISHES, FAMILY 
GOBIESOCIDAE, WITH DESCRIPTIONS OF NEW GEN- 
ERA AND FORMS 



By Leonard P. Schultz 



Recently, while attempting to identify some specimens of cling- 
fishes from the fresh waters of Venezuela and Colombia, South 
America, it became clear to me that the American Gobiesocidae were 
in a state of confusion greater than I had suspected from previous 
work on the group. A search of the literature did not reveal any 
attempt to strai<i:hten out the classification or nomenclature of this 
family. Mew species have been described with regularity, but in 
many cases the doscribers have ignored the same species long ago 
named from nearly the same localities. I have made no attempt in 
this revision to record all the miscellaneous references to American 
Gobiesocidae, but I have included the most important contributions. 
To record every reference in the literature would require one to 
examine the specimens in most of the important museums of the 
worM, which is not possible at the present time. However, that will 
eventually iiave to be done if the identifi«ations recorded in numerous 
instances are to be corrected. 

It was found necessary to dissect tlic skin away from the front of 
all median fins in order to count all the fin rays. Vei*y few authors 
lia\t' counted tlie first one or two rays at the beginning of both anal 
and doisal fins. The shoi-t stubby ray on the dorsal edge of I he 
pectoral fin also is included in my counts. (See table L) 

The coloration is variable and cannot be used to separate species 
except to a limited extent. 

47 
G04148 — «4 1 



48 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

In this study of the American Gobiesocidae, 9 genera and 25 species 
and subspecies are recognized. By far the greatest majority of these 
occur in the tropical Pacific. Although the Gobiesocidae are mostly 
marine fishes, a few species have been taken in fresh-water streams 
a short distance above brackish water, clinging to the stones. 

Rimicola of the Pacific is represented in the Atlantic by the genus 
Acyrtus^ but the genus Infratridens has no known Atlantic counter- 
part. Arhaciosa has five forms in the Pacific and but one in the 
Atlantic. Sicyases occurs in the Pacific only at the southern part 
of South America. Cotylis has four species in the Pacific and one 
in the Atlantic. Sicyogaster has an interrupted distribution, with 
one species along the Pacific coast of southern South America and 
another along the west coast of the United States and British Colum- 
bia. Arcos has one species on each side of Central America. 
Gohiesox has two species in the Atlantic, and three in the Pacific 
confined to tropical waters. 

Cocos Island presents a problem that needs further study. So far, 
the Pacific representatives of Gohiesox cephalus, and Cotylis nigripin- 
nis from the Atlantic, have been taken only on Cocos Island. They 
are Gohiesox fulvus and Cotylis nigripinnis woodsi. 

The records of Gohiesox adustus (Pellegrin, Bull. Mus. Paris, vol. 7, 
p. 206, 1901, and Giinther, Biologia Centrali-Americana, Pisces, p. 4, 
1906) in the Rio Chapalagana at Tepico in the Rio Grande de San- 
tiago may be some other species. No description is given, and thus 
it is not possible to place the above record with any species until the 
specimens have been re-examined. 

While this study was being made, Dr. S. F. Hildebrand kindly 
turned over to me some notes made by Dr. W. H. Longley on types in 
certain museums of Europe. Though most difficult to read, these 
proved of considerable value, and it was a pleasure to note that I 
came to the same conclusions that the late Dr. Longley had inde- 
pendently arrived at in regard to referring certain species to the 
synonymy of others. Although his notes were never published, some 
of the conclusions appeared during 1933 and 1934. 

The following key was prepared after examining the American 
clingfishes in the collections of the United States National Museum, 
as well as specimens lent by the Chicago Natural History Museum 
(F. M. N. H.) through the courtesy of Dr. K. P. Schmidt and Mrs. 
Marion Grey, and others lent by the University of Michigan Museum 
of Zoology through the courtesy of Dr. Carl L. Hubbs. Dr. C. M. 
Breder, Jr., of the American Museum of Natural History, kindly 
allowed me to examine the holotype of Gohiesox yuma Nichols. 



REVISION OF AMERICAN CLINGFISHES — SCHULTZ 49 

KEY TO THE GENERA AND SPECIES OF AMERICAN GOBIESOCIDAE 

1'/. Groove bt'twivn tip of snout iiiul upiht lip of prt'iuaxilluries extonding around 
front of snout and not forming a convex curve dorsally over tip of snout; 
width of middle of upper lip narrow, about the same as hiterally, and ap- 
proximately equal to width of pupil; gill membranes attached opposite 
tliird to fifth upper pectoral fin rays; axial flap of skin behind pectoral fin 
with its upi>er edge attached at midbase of pectoral fin or below midbase; 
fleshy pad on outer pectoral base present only ventrally, without a free 
margin posteriorly and enlarged or swollen at lower posterior corner of 
l)ectoral fin base: lower first to fifth pectoral raj's short, about half length 
of longest pectoral ray, eighth and ninth much longer than lower pectoral 
rays; anal rays G to 8; dorsal rays G or 7 (all rudiments counted as 
one ray). 
2rt. Incisorlike teeth at front of lower jaw with 4 minute points,^ these at front 
of upper jaw mostly conical ; each jaw with 1 or 2 inner rows of minute 
conical teeth ; axial fiap of skin behind pectoral fin attached at lower part 
of pectoral fin base; anal origin a little behind a vertical line through 
dorsal origin ; greatest depth of body 5V_> to 6Vj, length of head 3 to 3%. 
greatest width of head 4i/j to 5, length of disk 5 to 5Vj, all in standard 
length ; length of disk about equal to distance from tip of snout to front 
of disk ; pectoral rays about 1!) to 21 ; color when alive green or reddish, 
with or without light spots {Acyrtus, new genus) (Florida Keys and 

West Indies) Acyrtus rubiginosus (Poey) 

26. Incisorlike teeth at front of lower jaw with smooth tips ; middle front teeth 
of upper jaw conical ; teeth in inner rows of both jaws shorter, smaller, 
and conical : axial fiap of skin behind pectoral fin attached opposite middle 
of pectoral base; greatest depth of body 8 or 9, length of head 3Vj to 3%, 
greatest wiilth of head 5, length of disk 5Vo, all in standard length ; anal 
origin a little in advance of dorsal origin ; interorbital space 3^/^ in head, 
eye 1V> in interorbital space; length of disk about equal to caudal 
I)eduncle; lower pectoral rays shorter, second and third from bottom about 
half length of longest pectoral fin rays ; pectoral fin rays about IG or 17 
(Rimicola Jordan and Evermann) (Todos Santos Bay, Baja California 
to Monterey Bay and west coast of Vancouver Island, British Colum- 
bia) Rimicola eigenmanni (Gilbert) 

lb. Tip of snout formed by premaxillaries, which are much wider at middle of 

snout than laterally, groove arched dorsally over tip of snout ; axial Hap of 

skin behind p<'Ctoral fin with its upper etlge attached much above midbase of 

this fin ; lower first to seventh pectoral fin rays not shortened, about as long 

as eighth or ninth from bottonj. 

3fl. Anterior twth of lower Jaw trlfid incisors, trifid tips usually evident, except 

middle 2 or 3 sometimes worn off smooth although 1 or 2 of more laterally 

placed incisors at front of lower jaw always trifid. 

An. CJlll membrane attached opjiosite third to fifth jx-cloral lln rays; front 

teeth of upper Jaw smooth liiiped incl.sors (sometimes fiattened-cor)l- 

fonn) ; iront of both Jaws with 1 or 2 inner rows of small conical teeth 

behind outer row of enlarged Incl.sorlike teeth, sometimes these inner 

rows apparently represented by oidy 2 or 3 teeth ; fleshy pad on outer 

base of pectoral fin with free posterior margin ending a little below 

attachment of gill membraneH ; greatest width of head 3, length of head 



' SoiiiPtiiiKfl the middle two tpctli arc worn down n<'nily HnirMdli. jih In tlio type of '/. 
hrrtillinun Illldclirand and OlnxliiirK. 



50 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 93 

2% to 2%, greatest depth of body 5 to 6, length of disk 31/2, all in stand- 
ard length ; length of disk much greater than distance from tip of snout 
to front of disk ; distance from dorsal origin to midbase of caudal fin 
contained 1% to 1% times in snout tip to dorsal origin; anal origin 
under base of the third or fourth dorsal fin ray ; caudal peduncle short, 
its depth about equal to its length and about 3 times in base of dorsal 
fin; dorsal fin rays 11 to 13, anal 10 or 11, pectoral 18 to 21 (usually 
19 or 20) {Infratridens, new genus) (Gulf of California; southern 

California) Infratridens rhessodon (Rosa Smith) 

4&. Gill membranes joined opposite upper edge of pectoral fin base ; incisorlike 
teeth of both jaws with trifid tips, except middle pair or two sometimes 
smooth-tipped ; teeth in both jaws in a single row, lateral 2 to 4 conical 
and last 1 or 2 sometimes strong canines ; outer lower base of pectoral 
fin with fleshy pad poorly developed and without any trace of a free 
margin ; pelvic fins joined about halfway out fourth to sixth pectoral 
fin rays and not near base ; dermal flap in axile of pectoral fin joins oppo- 
site fourth to tenth pectoral fin ray ; width of head BV2 to 6, length of 
head 2% to 5, greatest depth of body 6 to 10 (except in eos), length of 
disk 4 to 6 (except in eos), all in standard length; opercular spine 
not strongly developed and not reaching to rear of head {Arhaciosa 
Jordan and Evermann). 
5a. A pair of black spots (more or less ocellate) on back behind head over 
pectorals usually distinct, each spot well separated ; dorsal surface of 
back in front of dorsal origin variously barred or mottled or dark 
spotted but without 3 hourglass-shaped large dark blotches. ( Species 
inhabiting waters of the Pacific coast and offshore islands.) 
6a. Distance from base of last dorsal ray to midcaudal fin base con- 
tained l%o to 1% times in length of dorsal fin base; least depth 
of caudal peduncle about 1.0 to 1% times in length of caudal 
peduncle (from base of last anal ray to midcaudal fin base) ; inter- 
orbital space longer than length of ^out ; dermal flap of skin 
in axis of pectoral fin with its upper edge joined to pectoral fin 
base opposite fifth to eleventh pectoral ray; dorsal rays 10 or 11 
(usually 10), anal rays 7 to 9 (usually 8 or 9) ; pectoral fin rays 
19 or 20 ; middle teeth of both jaws with trifid tips, middle denticle 
usually longest on lateral teeth, worn down in adults (Gulf 

of California) Arbaciosa humeralis (Gilbert) 

6b. Distance from base of last dorsal ray to midcaudal fin base con- 
tained 0.75 to 0.9 in length of dorsal fin base ; least depth of caudal 
peduncle 1.6 to 2 times in length of caudal peduncle. 
7a. Pectoral fin rays 22 to 24 (usually 23) ; dorsal rays usually 8 (7 
to 9), anal 7 or 8 (usually 8) ; greatest width of head contained 
Sy2, length of head 2% to 2%, in standard length ; snout a little 
longer than width of interorbital space; middle teeth of both 
jaws with trifid tips (Mazatlan, Mexico). 

Arbaciosa eos (Jordan and Gilbert) 

7b. Pectoral fin rays usually 19 to 21 (rarely 22). 

8a. Middle incisorlike teeth of both jaws (at least on adults) with 

smooth tips, the lateral incisors trifid ; pectoral fin rays usually 

about 21 ; bony ridges on snout weakly developed. 

9a. Dorsal rays 8 to 10 (usually 9) ; anal 8 or 9 (usually 8) 

(Peru) Arbaciosa pyrrhocincla pyrrhocincla (Cope) 

9b. Dorsal rays 8 ; anal 7 or 8 (Galapagos Islands). 

Arbaciosa pyrrhocincla truncata Heller and Snodgrass 



REVISIOX OF AMERICAN CLINGFISHES — SCHULTZ 51 

8b. Middle Incisorlike teeth of both jaws usually trifid, seldom worn 
ciCf smooth even on adults; dorsal rays 6 to 8; anal G or 7; 
jiectoral 19 to 21 (usually IJ) or 20) ; bony ridges on upper 
part of snout rather well developed (Ecuador to Gulf of 

California) Arbaciosa rhodospila ((Jiinther) 

56. Three or four larixe hourglass-shaped daric brown or blackish blotches 
from in front of dorsal fin to rear of head ; a fainter one sometimes 
on top of head ; side of head with four oblique bars and sides of body 
with dark bars ; incisors with trifid tips; dorsal rays 7 to 9; anal 6 to 
9 (rarely G or 9) ; pectoral 18 to 23 (West Indies; Guatemala to 

Brazil) Arbaciosa fasciata (Peters) 

36. None of the teeth with trifid tips. 

10a. Middle pair of incisors on both jaws much broader and longer than 

adjoining pairs ; posterolateral teeth small and conical ; rims of orbits 

bony, elevated ; opercular spine strongly developed and forming 

posteriormost tip of head ; valvular flap and margin of anterior nostril 

with its margin finely fringed with short cirri ; gill membrane attached 

at upper anterior edge of pectoral fin base; fleshy pad well developed 

on outer lower surface of pectoral base, with a free membranous 

edge posteriorly ending at ba.se of tenth to twelfth pectoral ray ; 

shoulder girdle with a free dermal flap extending dorsally nearly to 

attachment of gill membrane; anal ori^rin under base of second or 

third from last dorsal fin ray; disk large, its length about equal to 

head and contained about 2% to 2-i/f, in standard length ; anus just 

behind rear margin of disk; origin of dorsal fin a trifle closer to tip 

of opercular spine than midcaudal fin base; dorsal rays 10 or 11 

(usually 11) ; anal 8 or 9 ; pectoral 24 or 25 {Sici/ascH Miiller and 

Troschel ) . 

llo. Dorsal origin equidistant between midcaudal fin base and upper edge 

of gill opening to middle of length of upper pectoral rays; distance 

from base of last dorsal ray to midcaudal fin base in upper edge of 

gill opening to dorsal origin 1.80 to 2.35; length of caudal peduncle 

in snout tip to anal origin 15.50 to 7.36; base of dorsal fin in head 

1.90 to 2.40; ba.se of anal fin in bead 2.83 to 4.06; base of dorsal fin 

In snout tip to dorsal origin 3.70 to 4.G7 ; base of anal fin in snout 

tip to anal origin G.70 to 8.8.'i (Chile and I'eru). 

Sicyases sanguineus Miiller and Troschel 
116. Dorsal origin equidistant between iiiidcaudal fin base and middle 
of postorbital length of head ; base of last dorsal ray to midcaudal 
fin base In upper edge of gill opening to dorsal (trigin 1.(55; length 
of caudal iH?duncle in snout tip to anal origin 5.26; base of dorsal 
fln in head 1.55; base of anal fin in head 2.14; base of dorsal fin in 
snout tip to dorsal origin 3.04; iiasi' of anal fln in snout tip to anal 
origin 5.10 (Juan Ffrnrmdcz Island). 

Sicyases hildebrandi, new species 
106. Middle pair of Incisors not enlarged, all the Inci.sorllke or conical teeth 
at front of both Jaws of nearly same size and length; front of lower 
Jaw with small incisors In 2 or 3 pairs, with smooth tips; posterolat- 
eral teeth smaller, conical, sometimes one or two a little enlarged 
and almost cardnelike; ustuilly a small patch of very short conical 
teeth behind outer row of larger teeth at front of jaws b\it sometimes 
lacking fir reduced to 1 or 2 teeth ; rims of orbits not elevated or bony ; 
anterior nostril with a dermal flap, sometimes with liifid or even 



52 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

multifid tips arising on posterior rim, but nostrils not fringed witli 
short cirri. 
12a. Short blunt papillae on lips and around mouth generally, these in 
form of short barblets, arrangement as follows : Median part of chin 
and lower jaw with 2 or 3 rows of papillae, or chin anteriorly with 
a pair of low lobes in form of reversed parentheses [ ) ( ] and 
sometimes at their inner tips a pair of papillae (more or less fused 
with anterior lobes in nigripinnis and in pinniger) ; an inner row 
of barblets lateral to median lobes, one pair on each side ; lower lip 
at each side of median part of chin lobelike, sometimes bearing 2 
small papillae ; along inner edge of groove of lower jaw are 2 or 3 
large papillae or knobs on each side; upper lip with a median 
papilla or knob and 5 more on each side ; front edge of snout above 
groove without papillae but laterally 3 to 5 knobs or papillae 
present or absent; sometimes another papilla occurring behind 
rictus and still another below rictus; gill membranes joined oppo- 
site fifth to seventh upper rays of pectoral fin ; fleshy pad on outer 
base of pectoral fin with a free posterior membranous margin 
extending dor.sally to opposite attachment of gill membranes ; dorsal 
rays 10 to 19 ; pectoral fin rays 21 to 27 ; anus closer to anal origin 
than to rear margin of disk {Cotylis Miiller and Troschel). 
13a. Dorsal rays fewer than 15, counting all rudiments. 
14a. Dorsal rays 13 or 14; anal 10; upper lip with papillae. 

15a. Pectoral fin rays 22; papillae around mouth short and knob- 
like (Gulf of California) Cotylis papillifer (Gilbert) 

15&. Pectoral fin rays 24 to 27; papillae around moiith more numer- 
ous, better developed, barbellike; lobe of lower lip next to 
middle of chin with two barbels (pi. 1, A) (Panama Bay to 

Ecuador and northern Peru) Cotylis microspilus (Fowler) 

14&. Dorsal rays 10 to 12 ; anal 8 to 10 ; papillae on upper lip, lobelike ; 
lobe of lower lip next to middle of chin without barbels (pi. 1, 
B) ; dorsal origin equidistant between midbase of caudal fin 
and middle of postorbital length of head to equidistant be- 
tween midcaudal base and upper base of pectoral fin ; color 
pattern variable ; median fins mottled, barred, or blackish with 
tips of rays white. 
16a. Depth 41/2 to 61/2 ; eye 3.1 to 3.6 in length of base of dorsal fin ; 
dorsal rays usually 11, anal usually 9, pectoral 22 to 26 
(Maryland to West Indies to Brazil). 

Cotylis nigripinnis nigripinnis Peters 
166. Depth about 6% or 7; eye 2.8 in length of dorsal fin base; 
dorsal rays 10, anal 8, pectoral 22 (Cocos Island). 

Cotylis nigripinnis woodsi, new subspecies 

13&. Dorsal rays 17 to 19 (counting all rudiments) ; anal 10; anal 

origin under base of ninth or tenth dorsal fin ray or under middle 

of base of dorsal fin ; origin of dorsal fin a little closer to tip of 

snout than midbase of caudal fin ; papillae on upper lip knoblike 

(Gulf of California) Cotylis pinniger (Gilbert) 

12&. No papilla on upper lip, lobelike structures occurring around lips of 
lower jaw when best developed being low knobs or ridges, chin 
lacking inner series of papillae as described for Cotylis. 
17a. Gill membranes joined at upper edge of pectoral fin base, some- 
times a little anteriorly, giving appearance of being opposite 



REVISION' OF AMERICAN CLINGFISIIES — SCIIULTZ 53 

bases of uppor flrst to third pectoral fin rnys or the orbits larper 

than iiiterorbifal space; liioisorlike teeth at front of lower jaw 

projeotinp forward in a nearly horizontal or oblique direction, 

middle pair a little larger than those laterally. 

18o. Anal rays 10 to 14; dorsal 12 to 16, pectoral 10 to 23 (counting 

all rudiments) ; fleshy pad on outer margin of pectoral fin 

base very well developed and free membranous border along 

its posterior edge extending up to or beyond twelfth pectoral 

ray from dorsal edge; interorbital space equal to or wider than 

eye; least depth of caudal peduncle 4^^ to 5Vi times in dorsal 

origin to midcaudal tin base ; anal origin under anterior third of 

dorsal fln base {Sicyof/ai^tcr Brisout de Barneville). 

19a. Anal rays 10 or 11 ; dorsal 12 or 13 ; free margin of fleshy pad 

on pectoral fin base ending abruptly opposite ninth to twelfth 

ray from upper edge of pectoral fin base; eye 1 to l^^ In 

interorbital space and 4 or 5 in head ; anus a little closer 

to i-ear margin of disk than to anus; origin of dors&l fln 

equidistant between midbase of caudal fin and anterior half 

of postorbltal length of head (rem and Chile). 

Sicyog-aster marmoratus (Jenyns) 
196. Anal rays 12 to 14 ; dorsal 13 to 16 ; fleshy pad on outer base 
of pectoral fin with free posterior margin ending gradually, 
about opposite flrst to third upper pectoral fin ray ; eye 1% 
to 2 in interorbital space; length of disk about 3, head about 
214. greatest depth 4% to 5, all in standard length ; anus 
much closer to anal origin than rear margin of disk; origin 
of dorsal equidistant between midbase of caudal fln and 
rear of head; interorbital space about equal to snout (San 
Diego to Queen Charlotte Islands, British Columbia ; I'uget 

Sound) Sicyogaster maeandrlcus (Girard) 

186. Anal rays 7 or 8; dorsal 7 to 9; pectoral 22 to 25; diameter of 
eyes greater than interorbital space, the latter about % to % 
in eye; color usually reddish when alive (Arco/<, new genus). 
20a. Free margin of fleshy pad on pectoral fln base ending op- 
posite thirteenth ray from uj^per edge of pectoral fin base; 
Interorltital space G or 7 in head; least depth of caudal 
peduncle 3Vi' times in distance from midcaudal fin base to 
dcrsal origin and 1 '/•• in Its length; iielvi<-s fastened nearer 
base of r)ectoral rays than one-third way out (Oalapii^os 
Islands; Panama Bay; and MazatlAn, Gulf of California). 

Arcos poecilophthalmus (Jenyns) 

20^. Free margin of fiesby pad on pectoral fin liase ending opi)oslte 

sixteenth to nineteenth r;iy from upiK-r edge of pectoral fln 

base; interorbital ~> or <i limes In bead; least deptii of caudal 

peduncle 4 times in flistance from midcaudal base to dorsal 

origin and r4ft In its b-ngth; pelvics fastened about on<'-thlrd 

way out lower i)ectoral rays (Bahama Islands; West Indies). 

Arcos macrophthalmua (G(lnther) 

176. 0111 membranes Joined opposite third to seventh upp<'r pectoral fln 

rays .somewhat more anteriorly than In Cotyliit; liiclsorllke teeth 

at front of lower Jaw not prnj»H'tlng horizontally forward but 

curved oblifpiely upward so as to nearly opjio.se those In upper 

Jaw, the pair of incisors at middle of lower Jaw nearly same size 

as a«lJoinlng ones; outer surface of p<'<Moral fln base with a dls- 



54 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

tinctly fleshy pad, posterior margin free and joined opposite at- 
tachment of gill membranes {Gobiesox Lacepede). 
21ffi. Disk much greater than distance from tip of chin to front of disk. 
22a. Origin of dorsal fin equidistant between midcaudal fin base and 
rear one-third of pectoral fin rays or a little behind them ; anal 
origin under fifth dorsal fin ray, behind middle of base of rays 
of that fin; teeth of lower jaw not projecting forward in a 
nearly horizontal position but directed nearly straight up- 
ward in adults, a little more oblique in young specimens; 
head 2.2 to 2.7, disk 2.6 to 3.3, and depth 4 to 5.5, all in 
standard length ; dorsal rays 8 or 9, anal 5 to 7, i)ectoral 18 
to 21 ; anus equidistant between anal origin and rear margin 
of disk or a little nearer to anal origin ; eye 1% (young) to 5 
(adults) times in interorbital space. 
23a. Length of disk when measured from its rear margin reaches 
nearly to end of anal fin usually from midbase to base of 
last anal ray ; small dark spot often present near front of 
base of dorsal fin (Costa Rica, West Indies, to Brazil). 

Gobiesox cephalus Lacepede 

23&. Length of disk when measured from its rear margin reaches 

only to base of first or second anal fin ray ; front of dorsal 

with a large dark blotch not at base of fin (Cocos Island). 

Gobiesox fulvus Meek 

226. Dorsal origin equidistant between midcaudal fin base and upper 

base of pectoral fin or rear of head ; dorsal rays 10 or 11, anal 

8, pectoral 19 to 21 ; gill membrane attached opposite fourth 

to seventh upper pectoral fin rays ; interorbital 3% to 4i/^ in 

head ; distance from base of last dorsal ray to midcaudal base 

2% to Sli times in distance from dorsal origin to rear of head ; 

anus equidistant between or closer to anal origin than rear 

margin of disk ; fleshy pad on pectoral fin base with posterior 

margin free all way up to attachment of gill membrane ; anal 

origin under fifth dorsal ray ; depth of caudal peduncle equals 

its length. 

24a. Eye IV2 to 2 in interorbital space (Texas; British Honduras; 

Bahamas and West Indies). Gobiesox punctulatus (Poey) 

246. Eye 0.9 to 1.1 in interorbital space (Pacific-Mazatlan). 

Gobiesox adustus Jordan and Gilbert 
216. Disk about equal to distance from tip of chin to front of disk; 
pelvic fins attached about one-third way out pectoral fin rays; 
length of disk equal to distance from rear margin of disk to 
anus or 1% times from disk to anal origin; head 2.9, disk 3.8 
to 4.2, depth 5 or 6, width of head 3%, all in standard length ; 
eye 1% to 2 in interorbital space; interorbital 3^ and disk 
1% in head ; dorsal origin equidistant from midcaudal fin base 
and middle of length of pectoral fin ; distance from last dorsal 
ray to midcaudal fin base 4 times in distance from rear of head 
to dorsal origin ; anus closer to anal origin than rear margin 
of disk ; least depth of caudal peduncle greater than length of 
caudal peduncle from base of last anal ray to midcaudal fin 
base; opercular spine not well developed; anal origin under 
base of about the sixth dorsal ray ; dorsal rays 11 or 12 ; anal 
7 or 8; pectoral 18 to 20 (Gulf of California). 

Gobiesox funebris Gilbert 



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56 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

ACYRTUS, new genus 

Genotype. — Sicyases ntbiginosus Poey. 

This new genus is characterized by the lack of a dorsal curve of the 
premaxillary groove along upper lip over the snout tip in conjunction 
with the incisorlike teeth of the lower jaw having four minute joints. 
It may be distinguished from all other genera of American Gobieso- 
cidae by the key on page 49. Other characters are those of the 
genotype. 

Named Acyrtus in reference to the absence of the dorsal curve of the 
premaxillary groove over front of snout so common in all other Ameri- 
can clingfishes except Bimicola, to which this new genus is most closely 
related. 

ACYRTUS RUBIGINOSUS (Poey) 

Sicyases rubiginosus Poet, Synopsis piscium Cubensium, p. 391, 1868 (Palmasola, 

Cuba); Enumeratio piscium Cubensium, p. 124, 1876 (Cuba). — Jordan, 

EvEBMANN, and Clabk, Rep. U. S. Comm.. Fish, for 1928, pt. 2, p. 490, 1930 

(Matanzas, Cuba). 
Sicyases carneus Poey, Synopsis piscium Cubensium, p. 392, 1868 (Palmasola, 

Cuba) ; Enumeratio piscium Cubensium, p. 124, 1876 (Cuba). — Joed an, 

EvERMANN, and Clark, Rep U. S. Comm. Fish, for 1928, pt. 2, p. 490, 1930 

(Matanzas, Cuba). 
Sicyases carneus Voey—S. rubiginosus Poey, Longley, Carnegie Inst. Washington 

Year Book, No. 33, p. 271, 1934. 
fArbaciosa sp. Beebe and Tee-Van, Zoologica, vol. 10, No. 1, p. 252, 1928 (Lamentin 

Reef, Port-au-Prince Bay, Haiti). 
Gobiesox rubiginosus Jordan and Evermann, Rep. U. S. Comm. Fish and Fish. 

for 1895, App., p. 492, 1896 (Matanzas, Cuba) ; U. S. Nat. Mus. Bull. 47, pt. 3, 

p. 2337, 1898 (Cuba). 
Oobiesox carneus Jordan and Evermann, Rep. U. S. Comm. Fish and Fish, for 

1895, App., p. 492, 1896 (Matanzas, Cuba) ; U. S. Nat. Mus. Bull. 47, pt. 3, p. 

2337, 1898 (Matanzas). 
Gobiesox (Rimicola) beryllimis Hildeibrand and Ginsbttbg, Bull. U. S. Bur. Fish., 

vol. 42, p. 213, fig. 5, 1927 (Key West, Boca Chica, Fla. ) . 
Rimicola beryllinus Jordan, Evermann, and Clark, Rep. U. S. Comm. F^sh. for 

1928, pt. 2, p. 490, 1930 (Key West). 

Remarks. — An examination of the type of hei^yllinus indicates that 
the incisorlike teeth at the front of the lower jaw have four minute 
points, although the middle ones are worn down a little. The teeth 
along with other characters in the description of heryUmus are in need 
of rechecking, but the type appears to have been dried out sometime 
and is not in first-class condition. Hildebrand and Ginsburg's draw- 
ing, "figure 5," has the lower rays of the pectoral fin twice too long and 
the disk is not quite long enough. The fin rays were not correctly 
counted in the original description. 

Material examined.— Cvba : U.S.N.M. Nos. 82581 and 82582, totaling 
12 specimens. Florida : U.S.N.M. Nos. 87533 (holotype of beryllimis) 
and 116936, one specimen. 

Range. — Florida Keys and West Indies. , 



REVISION OF AMERICAN CLINGFISHES — SCIIULTZ 57 

Genus RIMICOLA Jordan and Evermann 

Rirnirola Jokuan and Evkkx[an.n, in Jordan, Pr«)<-. California Acad. Sci., vol. 6, 
p. 231, 1S96. (Genotype: Oobiesox muscarttm Moek and Plerson [ = Oobie80X 
eigenmanni Gilbert].) 

RIMICOLA EIGENMANNI (Gilbert) 

Gobicsox eigenmanni GiLBtJiT. Proc. U. S. Nat. Mus., vol. 13, p. 90, 1890 (Point 
Loma, near San Diego, Calif.). 

Oobiesox rhcssodon Rosa Smith, Proc. U. S. Nat. Mus., vol. 7, p. 553, 1885 (San 
Cri.stubal. P.aja California t. 

liimicola cifjenmanni Jokdan. Proc. California Acad. Sci., vol. 6, p. 231, pi. 32, 
1S9C. — Jordan and Evebmann, Rep. U. S. Conuu. Fish and Fish, for 1895, App., 
p. 4rt2, 189G (Point Loina) : U. S. Nat. Uus. Bull. 47, pt. 3, p. 2339, 1898 
(Point Loma, San Cri.«^tObal Bay).— Snyder, Proc. U. S. Nat. Mus., vol. 35, 
p. 183, 1908 (Todos Santos Bay, Baja California, northward to Pacific 
Grove, Calif.). — Jordan, Evekmann. and O.ark, Rep. U. S. Comm. Fish, for 
1928. pt. 2, p. 490, 1930 (Point Loma to Monterey).— Wiijjt, Copeia, 1936, 
No. 2, p. IIG (Round Island Flats, Clayoquot Sound, British Columbia). 

Rimicola muxraruyn Jordan, Proc. California Acad. Sci., vol. 6, p. 231, 1896. — 
Jordan and Evermann, Rep. U. S. Comm. Fish and Fish, for ISOn, App., 
p. 492, 1896 (Monterey Bay) ; U. S. Nat. Mas. Bull. 47, pt. 3. p. 2338, 1898 
(Monterey Bay). 

Oobiesox muscantm Meek and Pierson, Proc. California Acad. Sci., vol. 5, p. 571, 
pi. 71, 1895 (Monterey Bay, Calif.). 

Remarks. — My counts appear to disagree with those made by 
Snyder, Wilby, and other authors because I have inchided all the 
rudimentary fin rays at the beginning of each fin, apparently not 
counted previously. To be certain of my fin ray counts, the skin 
was di.s.sected away from one side of each fin at the base of the an- 
terior fin rays. This revealed usually one or sometijnes two rudi- 
mentar}' fin rays, and thus the counts by Snyder and by Wilby should 
be increa.sed by one or two to bring them into line with my counts. 

Material examined. — British Columiua (Vancouver Island) : 
U.S.X.M. No. 53800. California: U.S.N.M. Nos. 44372 (holotype 
of riqrnmnnni), 48875 (cotype of rnvsranivi) ., 49570, and G1055; 
U.M..M.Z. Xos. 64260, 64261, 642(52, 6364S, 63646, 63647, totaling 17 
specimens. 

Range. — Todo.s Santos Bay, Baja Califomia, to jMonterey Bay, and 
west coast of Vancouver Ishind. 

INFRATRIDENS, new genus 

Genotype. — Oobiesox rliessodon Rosa Smith. 

This new genus may be recognized from all otjier genera of Gobie- 
.socidae by the characters descriFn'd in the key on |)ag{'s 40-.50. It is 
di.stinguislu'd by its trifid incisorlikc lc(>th at front of lower jaw, the 
smooth-tipped teeth in upper jaw, and the convex premaxillury groove 
across front of snout. Otber characters are those of the genotype. 



58 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

Named hifratridens in reference to the trifid teeth at front of 
lower jaw. 

INFRATRIDENS RHESSODON (Rosa Smith) 

GoMesox rhessodon Rosa Smith, in Jordan and Gilbert, Proc. U. S. Nat. Mus., 
vol. 4, p. 63, 1881 (Point Loma) (nomen nudum). — Rosa Smith, Proc. U. S. 
Nat. Mus., vol. 4, p. 140, 1881 (San Diego, Calif.). 

Arbaciosa rhessodon Jordan, Proc. California Acad. Sci., vol. 6, p. 230, pi. 36, 
1896. — Jordan and Evermann, Rep. U. S. Comm. Fish and Fish, for 1895, 
App., p. 492, 1896 (San Diego; Gulf of California) ; U. S. Nat. Mus. Bull. 
47, pt. 3, p. 2340, 1898 (San Diego; Gulf of California).— Jordan, Evermann, 
and Clark, Rep. U. S. Comm. Fish, for 1928, pt. 2, p. 490, 1930 (San Diego; 
Gulf of California). 

Material examined. — Caufornia: U.S.N.M. Nos. 5246, 28396 (3 
types of rhessodon), 34765, 41975, 49574, 67312, 104193, 117642, total- 
ing 24 specnnens; U.M.M.Z. Nos. 63650, 63653, 63651, 63649, 63652, 
totaling 69 specimens. Catalina Island: U.S.N.M. No. 121964 and 
U.M.M.Z. No. 64263, totaling 8 specimens. Baja California: 
U.S.N.M. Nos. 36948 and 79149, totaling 2 specimens. 

Range. — Southern California to Baja California. 

Genus ARBACIOSA Jordan and Evermann 

Artaciosa Jordan and Evermann, in Jordan, Proc. California Acad. Sci., vol. 6, 
p. 230, 1896. (Genotype: OoUesox JiumeraUs Gilbert.) 

arbaciosa HUMERALIS (Gilbert) 

Oobiesox humeralis Gilbert, Proc. U. S. Nat. Mus.. vol. 13, p. 95, 1890 (Puerto 
Refugio, Angel Island). — Pellegrin, Bull. Mus. Hist. Nat. Paris, vol. 7, 
p. 162, 1901 (Gulf of California). 

Arbaciosa humeralis Jordan, Proc. California Acad. Sci., vol. 6, p. 230, pi. 35, 
1896. — Jordan and Evermann, Rep. U. S. Comm. Fish and Fish, for 1895, 
App., p. 491, 1896 (Gulf of California) ; U. S. Nat. Mus. Bull. 47, pt. 3, p. 
2341, 1898 (Angel Island; La Paz). — Jordan, E\t:rmann, and Clark, Rep. 
U. S. Comm. Fish, for 1928, pt. 2, p. 490, 1930 (Gulf of California). 

Material examined. — Gulf of California : U.S.N.M. Nos. 44374 
(cotype of humeralis), 125008 (cotype of humeralis), 46693, 48259 
(4 cotypes of humeralis), totaling 8 specimens; F.M.N.H. No. 3336, 
44 specimens ; U.M.M.Z. No. 136128, 1 specimen. 

Range. — Gulf of California. 

arbaciosa EOS (Jordan and Gilbert) 

Oobiesox eos Jordan and Gilbert, Proc. U. S. Nat. Mus., vol. 4, p. 360, 1882 
(Mazatlan). 

Arbaciosa eos Jordan, Proc. California Acad. Sci., vol. 6, p. 230, pi. 37, 1896 
(Mazatlan). — Jordan and Evermann, Rep. U. S. Comm. Fish and Fish, for 
1895, App., p. 491, 1896 (Pacific coast of Mexico) ; U. S. Nat. Mus. Bull. 47, 
pt. 3, p. 2343, 1898 (Mazatlan). — Jordan, Evermann, and Clark, Rep. U. S. 
Comm. Fish, for 1928, pt. 2, p. 491, 1930 (Pacific coast of Mexico). 

Material examined.— M-exico (Mazatlan) : U.S.N.M. No. 30889 (18 
cotypes of Gohiesox eos), C. H. Gilbert. 



REVISION OF AMERICAN CLINGFISHES — SCHULTZ 59 

Range. — Mazatlan, Mexico. 

ARBACIOSA PYRRHOCINCLA I'YUKHOCINCLA (Cope) 

Sicyases pyrrhorincUis Cope, Proc. Anier. Pliilos. Soc, vol. 17, p. 43, 1S77 (Peru). 
Arhaciosa pitrrhocinchis Abbott, Proc. Acad. Nat. Sci. Philadelphia, 1S99, p. 363 

(Peru. Pwasmayu Bay ?). — E\T':i£MAnn and R.vdcuite, U. S. Nat. Miis. 

Bull. 95, p. 155, 1917. 
Arhaciosa hicroglnphka Evp^imanx and Raucliffe, U. S. Nat. Mus. Bull. 9.', p. 155, 

pi. 14, fig. L', 1917 (Ldbo.s de Afuera, Porn). 
? Arbariofia petcrsii (nan Garniau) AiiBorr, Proc. Acad. Nat. Sci. Philadelphia, 

1S99, p. 363 (Peru) [not based on any .spociuicu]. 
? Oobicsox zebra (non Jordan and Gilbert) Rkoan, Ann. Mag. Nat. Hist., ser. 8, 

vol. 12, p. 280, 1933 (Lobos de Tierra, Peru). 

Material examined. — Peru (Lobos de Afuera Islands) : U.S.N.M. 
Nos. 77561 (type of hierogly phica) ^ 77565 (10 cotypes of hieroglyph- 
ica), 101703, 101704, 101705, 12-8175, totaling 21 spcciineii.s. Also from 
Peru : U.S.N.M. Nos. 88817, 88827, 88828, 119753, 128174, totaling 19 
specimens. 

Range. — Peru. 

ARBACIOSA PYRRHOCINCLA TRUNCATA Heller and Snodgrass 

Arhaciosa trunrata IIkixitk and Snodcr-vss, Proc. Washington Acad. Sci., vol. 5, p. 
210, pi. 14, liX)3 (Tagu.s Cove, Albemarle Island, Galdpagos). — Kendall and 
lUDCLUFE, Mem. Mu.s. Comp. Zool., vol. 35, No. 3, p. 160, 1912 (Chatham 
Island).— IlKKUK, Publ. Field Mus. Nat. Hist., zool. ser., vol. 21, p. 391, 1936 
(South Seymour Island ; Eden Island). 

Oohieaox zebra (non Jordan and Gilbert) Gilbkrt, Proc. U. S. Nat. Mus., vol. 13, p. 
452, 1890 (Duncan Island, Gal.lpagos). 

Material eu^amined. — Galatagos Islands: U.S.N.M. Nos. 65427, 
101702, 101710, 101715, 101716, 101718, 109419, 116202, 119337, totaling 
36 specimens; F.M.N.H. No.s. 25105-25215, 41301-41303, 41214, 41215, 
41641^1646, totaling 62 specimens. 

Range. — Galapagos Islands. 

ARBACIOSA RHODOSriLA (Gunther) 

Oobicsox rhoflosj)ilus G: nthkk, Proc. Zool. Soc. London, vol. 0, p. 25, 1864 (Pan- 
ama) ; Trans. Zool. Soc. lyindon, vol. 6, pp. 300. 445, 1S09 (Panama). — Jordan 
and KVKBMANN, Rep. U. S. Comm. Fish and Fish, for 18JK>, App., p. 492, 1896 
(Panama) ; U. 8. Nat. Mus. Bull. 47, pt. 3. p. 2335, 1898 (Panama).— Bou- 
i,KN(iKB. Bol. Mus. Zool. Anat. (."omp. Iniv, Torino, vol. 11, No. 335, p. 8, 1S99 
(Santa Kl«'na Bay, Iv-uador).— GiiJjarr and Stauks, Mem. California Acad. 
Sci., vol. 4, p. 1S9, 1904 (Panan)a).- Mkkk and IIildkiika.m), The marinr 
flslicH of Panama, pt. 3. p. 926, 192S (Panama and Santa EIrna Bay). — Joudan, 
EvKUMA.NN, and Cr.AiiK, Hep. U. S. Comm. Fish, for 192M. j.t. 2, p. 489, 1930 
(i'anama). 

Sicyases petersii Garman, Proc. Boston Soc. Nnt. Hist., vol. 18, p. 203, 1875 (San 
Joa6, San .Miffjifl, and Sabnga. all Pj-arl Island.s, Panama Bay). 

Oobicsox zebra Jokhan and Ghhfst, I'ror. U. S. Nnt. Mus., vol. 4, p. 359, 1882 
(Mnzatlfin). 



60 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

Arhaciosa sehra Jordan and Eveemann, Rep. U. S. Comm. Fish and Fish, for 1895, 
App., p. 403, 1896 (Mazatlan) ; U. S. Nat. Mus, Bull. 47, pt. 3, p. 2341, 1898, 
(MazatMn). — Mebsc and Hildebkand, Marine fishes of Panama, pt. 3, p. 927, 
1928 (Toboguilla Island). — Kendall and Radcliffe, Mem. Mus. Comp. Zool., 
vol. 35, No. 3, p. 160, 1912 (Toboguilla Island). — Jordan, Evermann, and 
Clabk, Rep. U. S. Comm. Fish, for 1928, pt. 2, p. 491, 1930 (MazatlSn.) 

Material examined. — Gui:f op CAiiiFORNiA : U.S.N.M. Nos. 29250 (47 
cotypes of seSra), 47496, 119716, 119717, 119754, totaling 56 specimens. 
Costa Eica: U.S.N.M. Nos. 92119 and 101711, totaling 3 specimens. 
Panama Bay: U.S.N.M. Nos. 65428 and 120435 (3 cotypes of petersii), 
totaling 8 specimens. Colombia : U.S.N.M. No. 101712, one specimen. 
Ecuador : U.S.N.M. Nos. 88826 and 101717, totaling 3 specimens. 

Range. — Gulf of California to Ecuador. 

ARBACIOSA FASCIATA (Peters) 

Sicyases fasciatus Peters, Monatsb. Akad. Wiss. Berlin, 1859, p. 412, May 8, 1860 
(Puerto Cabello [probably Venezuela]). — Gunthek, Catalogue of the fishes 
in the British Museum, vol. 3, p. 497, 1861 (Puerto Cabello) ; Trans. Zool. 
Soc. London, vol. 6, p. 390, 1869 (Puerto Cabello). — Guitel, Arch. Zo-ol. Exper., 
vol. 5, No. 5, pp. 645-652, 1906 (anatomy). — Jordan, Evermann, and Clark, 
Rep. U. S. Comm. Fish, for 1928, pt. 2, p. 490, 1930 (Puerto Cabello). 

Oohiesow rupestris Poey, Memorias sobre la historia natural de la isla de Cuba, 
vol. 2, p. 283, pi. 18, fig. 6, July 1860. 

Oobiesox fasciatus Jordan and Evermann, Rep. U. S. Comm. Fish and Fish, for 
1895, App., p. 492, 1896 (Puerto Cabello) ; U. S. Nat. Mus. Bull. 47, pt. 3, p. 
2338, 1898 (Puerto Cabello). 

Arhaciosa rupestris Jordan and Evermann, Rep. U. S. Comm. Fish and Fish, for 
1895, App., p. 492, 1896 (Cuba) ; U. S. Nat. Mus. Bull. 47, pt. 3, p. 2341, 1898 
(Cuba). — Beebe and Tee- Van, Zoologica, vol. 10, No. 1, p. 252, fig., 1928 
(Port-au-Prince Bay, Haiti). — Jordan, Evermann, and Clark, Rep. U. S. 
Comm. Fish, for 1928, pt. 2, p. 491, 1930 ( Cuba ) .—Parr, Bull. Bingham 
Oceanogr. Coll., vol. 3, art. 4, p. 136, 1930 (Green Cay, Bahamas). 

Sicyases rupestris Poey, Synopsis piscium Cubensium, p. 391, 1868 (Cuba) ; 
Enumeratio piscium Cubensium, pt. 2, p. 124, 1876 (Cuba). 

Arhaciosa minuta Meek and Hxldebrand, The marine fishes of Panama, pt. 3, 
p. 928, pi. 92, 1928 (Colon, Panama). 

Material examined. — St. Croix Island: U.S.N.M. Nos. 15382 and 
15431, totaling 13 specimens. Barbados : U.S.N.M. No. 86752, 1 speci- 
men. Cuba : U.S.N.M. Nos. 37414, 37421, 82580, totaling 15 specimens. 
Guatemala: U.M.M.Z. Field No. H35-138a, 6 specimens. Panama: 
U.S.N.M. No. 81523 (type of minuta). Brazil: U.S.N.M. Nos. 87799, 
87800-87803, 88042, totaling 19 specimens. 

Ra/nge. — ^West Indies ; Guatemala to Brazil. 

Genus SICYASES Muller and Troschel 

Sicyases Muxlee and Teosohel, in Miiller, Arch, fiir Naturg. (Wiegmaun), 9th 
year, vol. 1, pp. 297, 298, 1843 (genotype: Sicyases sanguineus Miiller and 
Troschel) ; Ber. Verb, preuss. Akad. Wiss., 1843, p. 212 (genotype: Sicyases 
sanguineus Miiller and Troschel) (ref. copied) ; Horae ichthyologicae. Be- 
schreibung und abbildung neuer Fische, pt. 3, p. 19, 1849. 



REVISION OF AMERICAN' CLINGFISHES — SCHULTZ 61 

Tomicodon Brisout de Barnb\jlle, Rpv. Zool. Soc. Cuv., vol. 9, p. 144, 1846 {Tomi- 
codun chilen^fis Brisout de Barueville) ; Echo Monde Savant, vol. 13, p. 535, 
184G. 

SICYASES SANGUINEUS MUIIer and Troschel 

Sicya^cs sanguineus Mi;ixt3 and Troschfx, in Miiller, Arch, filr Naturg. (Wieg- 
mann), 9th year. vol. 1, p. 298, 1843 (Chile) ; Horae ichthyologlcae, pt. 3, p. 19, 
pi. 3, fig. 1, 1849 (Chile). — e}i3NTHKK, Catalogue of the fishes in the British 
Mu.seuni, vol. 3, p. 404, ISOl (Chile: Valparaiso). — Delfin, CatAlogo de los 
peces de Chile, Valparaiso, p. 90, llMll (Bahia de Concepci6n ; Cavancha; 
Isla de Juan Fcrnaiulez ; Toni6; Talcahuaiio). 

Totnicodon chilcnxis BitisouT de Bauneaille, Rev. Zool. Soc. Cuv., vol. 0, p. 144, 
1846 (Valparaiso). 

? Oobicsox brcvirostris Gay, Ilistoria flsica y polilica de Chile . . . Zoologia, 
vol. 2, p. 335, pi. 9, fig. 1, 1848 (ref. copied). 

Gobiesox sanguineus Abbott, Proc. Acad. Nat. Sci. Philadelphia, 1899, p. 363, 
(coasts of Peru and Chile). — E\-ebuann and Radci.iffe, U. S. Nat. Mus. 
Bull. 95. p. 153. 1917 (Peru and Chile). 

Sicj/ases chilensis GI'-nther, CataloRuo of the fishes in the British Museum, vol. 
3, p. 497. ISGl (Valpar.ii.xo). — Delfin. CatAlogo de los peces de Chile, Val- 
paraiso, p. 90, 1901 (Valparaiso). 

Material examined.— Perv: U.S.N.M. Nos. 44130, 77512, 83029, 
91557. totalino: 11 .specimens. Chile (Valparaiso) : U.S.N.M. No. 
121950 and F.M.N.H. No. 32994, 1 specimen each number. 

Raiifje. — Peru and Chile. 

SICYASES HILDEBRANDI. new species 

Holotype. — U.S.N.M. No. 88818, the only known specimen, 69 mm. 
in stan(hinl lenfrth, collected by Dr. W. L. Schmitt at Cumberland Bay, 
Juan Fernandez Island, off Chile, December 1920. 

Description. — Certain measurements were made on the holotj'pe, 
and these alon^r with others made on three specimens of S. sanguineus 
are recorded in table 2. 

The following counts were made on the holotype: Dorsal rays 11; 
anal rays 9; pectoral rays 25-25; free edge of pectoral pad ends oppo- 
site 11-11 pectoral raj's counting down from the dorsal edge; gill 
membranes attached opposite upper edge of pectoral fin base. 

Head about 31/0, depth 0.9, di.sk 2.9, all in .standard length; eye V^/o 
in intt'rorbital space, the latter 2% in head (to upper edge of gill 
opening); dorsal origin equidistant between niidcaudal fin base and 
middle of postorbital length of head; anal origin under bases of 
second and third from last dorsal (in rays; tip of snout to tip of 
opercular spine equal to distance from upper edge of gill opening to 
dorsal origin; base of last dor.sal ray to niidcaudal fin ba.se V/^ in 
upper edge gill opening to dorsal origin; ba.se of dorsal fin l"/,o and 
l>ase of anal fin 2.1, lK)th in gill oj)eMing to dor.sal origin; ;iniis just 
hcliincl (lie icar niai-;:iii of the disk: lice iioslcrior miiiiriii of the 



62 



PROCEEDINGS OF THE NATIONAL MUSEUM 



fleshy pad on lower pectoral base ending opposite the eleventh ray 
from dorsal edge of pectoral fin; the middle incisors of both jaws 
much wider than the adjoining pair and those on lower jaw longer, 
but the two middle pairs of upper jaw about the same length; the 
interorbital space slightly concave ; both nostrils close in front of eye, 
the anterior one with a small fringed flap arising on its posterior edge. 

Table 2. — Measurements oj the two species of Sicyases, in hundredths of the 

standard length 



hildebrandi] 



Characters 



Holotype 



U.S.N.M. Nos. 



77512 77512 77512 83029 



Standard length (in millimeters) 

Length of head to -upper edge of gill opening 

Length of head to tip of opercular spine 

Greatest depth of body 

Greatest width of head 

Least depth of caudal peduncle 

Length of caudal peduncle 

Length of snout 

Diameter of eye 

Width of bony interorbital 

Postorbital length of head to upper edge of gill open- 



mg 



Postorbital length of head to tip of opercular spine.. 

Length of disk 

Distance from base of last dorsal ray to midcaudal 

base 

Disk to anus 

Anus to anal origin 

Snout tip to dorsal origin 

Snout tip to anal origin -. 

Snout tip to center of anus. 

Length of base of dorsal fin 

Length of base of anal fin 

Longest ray of dorsal fin 

Longest ray of anal fin — 

Longest ray of caudal fin 

Longest ray of pectoral fin 

Tip of snout to disk 

Origin of dorsal to upper edge of gill opening 



69 

28.3 

33.3 

14.5 

27.5 

7.82 
13.8 
10.6 

7.54 
12.2 

14.9 
19.1 
35.6 

21.7 
3.91 
19.6 
59.4 
72.5 
52.6 
19.5 
14.2 
12.5 
11.7 
24.5 
14.5 
13.3 
36.0 



59 

32.2 

35.6 

14.9 

29.1 

8.30 
13.9 
11.7 

7.63 
12.2 

16.4 
20.2 
36.4 

20.7 
3.90 
23.4 
62.7 
76.3 
53.2 
17.0 
11.4 
12.2 
11.2 
23.6 
15.3 
14.2 
37.3 



78.5 
33.8 
37.6 
15.9 
29.9 

7.77 
10.8 
12.2 

8.02 
14.0 

16.7 
21.9 

38.8 

17.5 
4.20 
20.4 
66.2 
79.7 
58.1 
15.5 
9.30 
12.5 
11.5 



14.3 
14.3 
37.6 



90 

35.3 

38.3 

16.1 

33.9 

8.33 
11.1 
13.1 

7.56 
16.0 

17.2 
23.3 
40.6 

19.1 
3.67 
21.6 
68.4 
80.0 
57.2 
14.7 
10.0 
13.3 
11.7 
23.5 
16.7 
15.5 
38.6 



159 
37.1 

44.3 
21.7 
37.1 

7.86 
13.8 
13.8 

6.2S 
18.2 

20.4 
28.0 
45.9 

16 
0.94 
20.4 
69.2 
80.5 
59.2 
17.9 
9.12 
14.1 



22.0 
18.2 
15.4 
37.7 



Remarks. — The chief differences between this new species and 
Sicyases sanguineus are in the more anterior position of the dorsal fin 
and the length of the bases of the dorsal and anal fins. The following 
measurements indicate the amount of the above differences, first for 
the new species then for sanguineus : Distance from base of last dorsal 
ray to midcaudal fin base in upper edge of gill opening to dorsal origin 
1.65 and 1.80-2.35; length of caudal peduncle in tip of snout to anal 
origin 5.26 and 5.50-T.36 ; base of dorsal fin in head 1.55 and 1.90-2.40; 
base of anal fin in head 2.14 and 2.8'3^.06 ; base of dorsal fin in snout to 



REVISION OF AMERICAN CLIXGFISHES — SCHULTZ G3 

dorsal origin 3.04 and 3.70^.67; ba>e of anal fin in snout to :inal orij^in 
5.10 and C.70-8.tv3 ; dorsal origin equidistant between niidcaudal fin base 
and middle of postorbital length of head for hildebrandi, and upper 
edge of gill ojiening to middle of length of upper pectoral rays for 
sanguhu'us. In general, the new species appears to be a little slenderer 
than sang^uineus. Presumably, when an adequate series of hildebrandi 
is collecteil from the Juan Fernandez Islands and studied, this new 
species may be best treated as a subspecies of sanguineus. 

Named hildebrandi for Dr. Samuel F. Hildebrand, senior ichthy- 
ologist, United States Fish and "Wildlife Service, who while working 
up a monograph of the fishes of Peru noticed this new fish and sug- 
gested that I describe it. It is with great pleasure that I name this 
new species in his honor and in recognition of his numerous and 
valuable contributions in ichthyologj'. 

Genus COTYLIS MuUer and Troschel 

Cotylis MCllkk and Troscuel, in Miiller, Arch, fiir Naturg. ( Wiegmann) , 9tb year, 
vol. 1, p. 297, 1S43 (genotype: Cotylis nnda Miiller and Troschel=Lcpado- 
gaKternudux Bl<»ch and Schneider=Go?>/r.«(oj?/7.i/n/iH.s' Jordan and Evermann = 
Gobiesox nigripinnis Peters) {Cycloptcrus nudus Linnaeus not identified) ; 
Horae iehthyalogicae, pt. 3, p. IT, pi. 3, lig. 2, 1840 (genotype: Cotylis nuda 
Bloch and Schneider). [Cotylis (non Midler and Troschel) Giiulher, Cata- 
logue of the fishes in the British Museum, vol. 3, p. 498, 1861, hut restricted to 
Cotylis fimbriata Miiller and Troschel, 1849, from Red Sea, and not described 
In 1843. Cotylis (liinther has the substitute name Cotylichthys Jordan, Proc. 
Acad. Nat. Sci. Philadelphia, 1919, p. 341.] 

liryssctacres Jokdan and Evkkmann, ih Jordan, Proc. California Acad. Sci., vol. 
«;. p. 230, 189G (genotype: Gobiesox pinniycr Gilbert) ; U. S. Nat. Mus. Bull. 
47, pt. 3, p. 2328, 1898 (genotype: G. pin«jV/cr Gilbert). 

Caulistius Joroan and Evehman.n, Rep. U. S. Comrn. Fish and Fish, for 1895, App. 
p. 491, 1896. (Genotype: Gobiesox papillifer Gilbert.) 

Ilryssophilus Jordan and Evermann, U. S. Nat. Mus. Bull. 47, pt. 3, pp. 2329, 2330, 
1898. (Genotype: Gobiesox papillifer Gilljert.) 

COTYLIS PAPILLIFER (Gilbert) 

Gobiesox papillifer Gimiebt, Proc. U. S. Nat. Mus., vol. 13, p. 90, 1890 (Magdalena 
Bay, Lower (.'alifornla). — Jokuan and Evi:u.\iann, R(»p. U. S. Comni. Fish 
and Fish, for lS9rj, App.. p. 4!»1, 1S96 (Magdalena Bay) ; U. S. Nat. Mus. Bull. 
47, pt. 3, p. 2330, 1898 (Magdalena Bay). 

Caulistius papillifer Jobda.n, Eveicmann, and Ci.abk, Rep. U. S. Conini. Fish, for 
1928, pt. 2, p. 488, 1930 (Magdalena Bay). 

Material examined. — Baja California (Magdalena Bay) : U.S.N. M. 
No. 44.'J7G (type of papillifer), collected by the Albatross. 
Range. — Magdalena Bay, Baja California. 



64 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

COTYLIS MICROSPILUS (Fowler) 

Plate 1, A 

CauUstiiis microspilvs Fowlee, Proc. Acad. Nat. Sci. Philadelphia, 1916, p. 412, 
fig. 4 (Panama Bay). 

Description. — Head contained about 2%, disk 2.9 or 3, greatest 
depth of body 5 to 6, in standard length; eye 21/2 in interorbital 
space ; disk a little shorter than length of head ; tip of chin to front 
of disk about % to % length of disk ; distance from base of last dorsal 
fin ray to midbase of caudal fin contained nearly three times in base 
of dorsal fin; dorsal origin nearly an eye diameter closer to mid- 
base of caudal fin than to rear of orbit; depressed anal fin reaching 
a trifle past a line through caudal fin base, and depressed dorsal fin 
reaching to opposite caudal fin base; upper lip on premaxillary with 
a median papilla and five more on each side, but none posteriorly on 
upper lip; middle of snout with three short papillae or knobs, but 
edge of snout above groove without papillae anteriorly but about five 
well-developed ones laterally ; another papilla behind rictus and one 
on lower lip below rictus; median part of chin and lower jaw with 
three rows of papillae, the most anterior being a pair of low lobes, 
next a pair of papillae, and the inner row consisting of two pairs 
of papillae, with the outer pair posterior to the anterior pair ; lower 
lip at each side of median part of chin forming a small lobe bearing 
two small papillae ; three large papillae along the inner edge of the 
groove along edge of lower lip on each side ; preopercular spine well 
developed; three or four pairs of small incisorlike teeth at front of 
lower jaw in outer row; teeth at front of upper jaw nearly conical; 
lateral teeth of both jaws conical; a small patch of teeth behind outer 
teeth at front of both jaws; interorbital space flat; anterior nostril 
tubular with a short dermal flap, sometimes branched, arising at the 
posterior rim of this nostril; shoulder girdle with a fleshy lobe and 
a shallow groove along its lower edge separating it from the lower 
less fleshy lobe; base of pectoral fin with a fleshy lobe, the posterior 
and ventral margins free, this free margin beginning at point where 
gill membrane is fused opposite base of sixth or seventh pectoral 
ray from dorsal edge of that fin; upper edge of axial dermal flap 
behind pectoral fin is fused to base of fin opposite ninth or tenth ray 
from dorsal edge of pectoral fin ; pelvic fin attached to near base of 
pectoral fin rays ; lower rays of pectoral fin nearly as long as middle 
pectoral fin rays ; margins of disk and pelvic pads of disk all covered 
with low flattened papillae; anus much closer to anal origin than 
to rear margin of disk. 

Coloration. — In alcohol, pale brownish everywhere on dorsal sur- 
faces of head and anterior parts of body profusely brown-spotted, 
these spots small and rather close together ; tips of all rays of median 



U. S. NATIONAL MUSEUM 



PROCEEDINGS. VOL 96 PLATE t 





B 



A, IxjAcr side of head of Colylis microjpiluj (F*'owlcr), V. S. N. M. No. 1U7H2. H, l/)wtr 
side of head of Cotylii nigripinnis nif,ripinnis Peters, U. S. N. M. No. 58829. Drawn 
bv Mrs \;,„r M Awl. 



REVISION' OF AMERICAX CLINGFISHES — SCHULTZ 65 

fins white; basally tlio dorsal, anal, and caudal fins are dark brown; 
more or less obscure pale bar across base of caudal fin. 

Material exaviincd. — The following three specimens, all collected by 
Dr. W. L. Schmitt, form the basis of the foregoing redescription of 
this species: 

U.S.N.M. No. 88822, 26.5 mm., Salinas, Ecuador, September 15, 1926. 
U.S.N.M. No. S8823. 58 mm., Guayaquil. Ecuador, 1026. 
U.S.N.M. No. 107142, 62.6 mm., Paita, Peru, October 7, 1926. 

The three young specimens listed below, also collected by Dr. 
Schmitt, are referred to this species with uncertainty. They appear 
to be more robust than the adults. 

U.S.N.M. No. 101713, 2 specimens, 14.5 and 15 mm., Cupica Bay, Colombia, 
January 26, 1935. 

U.S.N.M. No. 101938, 1 specimen, 9.5 mm., Cupica Bay, Colombia, .Tanuary 
26, 1935. 

Range. — Panama Bay to northern Peru. 

COTYLIS NIGRIPINNIS NIGRIPINNIS Peterg 
Pl.\te 1, B 

Cotiilis nigrijjinnis Peteks, Monatsb. Akad. Wiss. Berlin, 1S59, p. 412, May 8, 
ISfX) (Puerto Cal)ello fprol)al)iy Venezuela]). 

Oobifsox nigripinnin GCntiieb, Catalogue of the fi.-^hes in the British Museum, 
vol. 3, p. 502, 1S61 (Puerto Caliello) ; Trans. Zool. Soc. London, vol. 6, p. 390, 
1869 (Puerto Cabello). — Jokdan and Evctmann, Rep. U. S. Comm. Fisli and 
Fish, for l.s:).5, App.. p. 4lil. 189(; (Puerto Cabello) ; U. S. Nat. Mus. Bull. 47, 
pt. 3, p. 2331. 1898 (Puerto Cabello).— Metzelaab, Bijd. Dierk. Feest. Num. 
70th Geboortedag van Dr. Max Weber, pt. 22, p. 140, 1922 (Caracas Bay). — 
.loBDAN, E\T,RMANN, and CiJVBK, Kei>. U. S. Comm. Fish, for 1928, pt. 2, p. 
488, 19.30 f Puerto Cabello). 

Oohieaox slrumosun Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 22, j). 121, 1870 
(Hilton Head, S. C). — Jobdan anil Evekmann, Rep. U. S. Comm. Fish and 
Fish, for 1895, App., p. 491, 1896 (Carolina to Florida) ; U. S. Nat. Mus. Bull. 
47. pt. 3, p. 2.3;}3, 1898 (Ililioii Uvnd, S. C. ; Indian Illvcr, Fla. ; Titusville).— 
Hii.DEBKA.ND and ScnKoEnKii, Bull. U. S. Bur. Fish., vol. 43, pt. 1, p. 3.39, 1928 
(Chesapeake Bay). — Jobi>an, Evf^mann. and Ci..\bk, Kep. U. S. Comu). Fish, 
for 1J>28, pt. 2, p. 489, 1930 (Maryland to Florida).— Ix)noi.ey and Hiu)E- 
BRAND, Sysrematip eatalogue of the fishes of Tortugas, Florida, p. 284, 1941 
(TorfuKas. Fla.). 

Lrf>adt>iiUHt(r nudun {non Linnaeus) Bi.ocH and Scuneideb, Syslema Ichthyo- 
loBiae. p. 2, 1801 (locality ?). 

CotyliH nudn {nun LlriiiacuH ) Mi li.ku and 'I'uosc iiKi., ITorai- i( hthyoloKJcae. pt. 3, 
p. 17. pi. 3. fig. 2. 1849 [West Indies]. 

Gohirnox nudiin Brihott de Babnevim.e, Rev. Znol. Soc. Cuv., vol. 9, \>. 141, ISIO 
(no locality). — f;ONTHra, Catalogue of the fishes In the British Museum, vol. 
3, p. 502, 1861 (West Indies; Island of Cordova) ; Trans. Zool. Soc. London, 
vol. 6, p. ,3JK), 1869 ((Pardon). 

OnhicHoi ff]/rinuif Jordan and Evkicmann, Rep. U. S. Conun. Fish and Fish, for 
1895. p. 491, 1896 (nomcn nudum) ', U. S. Nat. Mus. Bull. 47, pt. .3, p. 2.'m. 
1898 (West Indies) (I)a8ed on Oobiesox nudug \non Linnaeus] GUnllier). — 



66 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

Jordan, Evekmann, and Claek, Rep. U. S. Comm. Fish, for 1928, pt. 2, p. 489, 
1930 (West Indies). 

GoMesox virgatulus Goode and Bean, Proc. U. S. Nat. Mus., vol. 5, p. 236, 1882 
(Gulf of Mexico) {nomen nudum). — Jordan and Gilbert, Froc. U. S. Nat. 
Mus., vol. 5, p. 298, 1882 (Pensacola, Fla. ) .—Jordan, Proc. U. S. Nat. Mus. 
vol. 7, p. 149, 1884 (Egmont). — Jordan and Evermann, Rep. U. S. Comm. Fish 
and Fish, for 1895, App., p. 491, 1896 (Pensacola Bay to Charleston, S. C.) ; 
U. S. Nat. Mus. Bull. 47, pt. 3 p. 2333, 1898 (Pensacola Bay north to Charles- 
ton). — Smith, Fishes of North Carolina, p. 374, 1907 (Beaufort Havbor; 
Fort Macon; Charleston).— Jordan, Evermann, and Clark, Rep. U. S. Comm. 
Fish, for 1928, pt. 2, p. 489, 1930 (Pensacola Bay to Charleston). 

OoMesox sancti-martini Metzelaae, Report on the fishes collected by Dr. J. Boeke 
in the Dutch West Indies, 1904-1905, pt. 1, p. 151, fig. 48, 1919 (St. Martin, 
Simsonsbay Lagoon). — Jordan, Evermann, and Clark, Rep. U. S. Comm. 
Fish, for 1928, pt. 2, p. 490, 1930 (West Inaies). 

OoUesox bariattilus Starks, The fishes of the Stanford Expedition to Brazil, p. 
73, pi. 14, 1913 (Natal). — Ribeiko, Fauna Brasiliense . . . Peixes . . . 
Gobiesocidae, p. 2, 1915 (Lagoa em Natal). 

"GoMesox yuma Nlchols=[non] Gobiesox vittatus Metzelaar=[non] Gobiesox 
punctulattis Poey," Longley, Carnegie Inst. Washington Year Book No. 34, 
p. 284, 1935. 

"Gobiesox virgatulus Jordan and Gilbert=G. strumosus Cope," Longley, Carnegie 
Inst. Washington Year Book, No. 33, p. 270, 1934. 

"Gobiesox barbatulus Starks= Goftiesoa? gyrimis Jordan and Evermann=Go&iesoa? 
nigripinnis Peters," Longley, Carnegie Inst. Washington Year Book, No. 34, 
p. 284, 1935. 

Gobiesox yuma Nichols, Bull. Amer. Mus. Nat. Hist., vol. 37, No. 37, p. 876, fig. 1, 
1917 (Sanibel Light, Fla., vrest coast). — ? Beeder, Bull. Bingham Oceanogr. 
Coll., vol. 1, art. 1, p. 85, 1927 (Royal Islands, Bahamas). — Jordan, Ever- 
mann, and Claek, Rep. U. S. Comm. Pish, for 1928, pt. 2, p. 490, 1930 
(Florida). 

Remarhs. — Miiller and Troschel's description of Cotylis nuda 
(1849, pp. 17-18) leaves little doubt that their species is the same as 
the one recognized here as nigripinnis, since small barbels are said 
to occur around the mouth and the coloration is brownish with streaks 
of dark spots. In addition, fin rays are given as dorsal 12, anal 7. 

Wlien the form along the Atlantic coast from Chesapeake Bay to 
the east coast of Florida is studied in the minutest detail, it may be 
recognized as distinct from nigripinnis, but I have not thoroughly 
investigated the variation in the various localities from Maryland to 
Brazil. There are several names available for the races or subspecies 
that may be recognized. 

Dr. S. F. Hildebrand kindly turned over to me the notes made by 
Dr. W. H. Longley at Amsterdam on the type of Gobiesox sancti- 
martini Metzelaar. I quote: 

T. L. [total length] G9 mm. D. 12, A, 8, P. 23-24 including a stub above. 
Diameter of eye (orbit) 3.0 mm. Interorbital width 7.0 mm. Nasal cirri 
expanded, bilobed, vpithout fringe. Twenty-nine coarse cirri, becoming bulbous 
under pressure of the tissue behind them, along front of ventral disk in single 



REVISION OF AMERICAN CLINGFISHES — SCHULTZ 



67 



series. The fleshy border lateral to them only slightly crenulated before the 
anterior ray of the ventral fin. The lower angle of the pectoral moderately 
prouiinent, not cxserted. Opercular cleft extending upward to the base of Gth 
ray, the fold before the base of the fin complete, continuous with the fleshy 
border of the operculum. Anterior teeth little if any flattened, the lateral in 
the upper j;iw running in behind tiie front but not as regularly as in some. . . 

Table .3. — Counts and measurements made on species of Cotylis, expressed 
in hundredths of the standard length 



Characters 




microtpilut 




nigripmnia 
nigripinnis 


ni(;TipinnU 
woodii 


U.S.N.M. 
No. 88822 


U.S.N.M. 
No. 88823 


U.S.N.M. 
No. 107142 


U.S.N.M. 
No. 87752 


U.S.N.M. 
No. 87752 


F.M.N.H. 
No. 41974 


Standard length (in milli- 
metors) 


26.5 
41.5 
15.8 
32.0 
8.30 

9.44 
11.7 

7.55 
10.9 
26.4 
35.2 

9.44 
13.6 

10.9 
6.04 
61.1 
73.2 
64.9 

32.5 
21.1 
17.0 
15.1 
30.6 
15.8 

14.0 
24-26 
14 
10 

6-7 


58 

41.2 
18.3 
31.1 
9.32 

8.62 
12.2 

5.86 
14.7 
24.2 
34.5 

11.2 
13.0 

11.7 
4.14 
59.2 
70.7 
66.9 

32.4 
22.1 
15.0 
12.4 
26.7 
15.5 

12.9 
27-27 
14 
10 

7-7 


62.6 
41.0 
18.2 
35.1 
8.79 

8.03 
13.3 

5.59 
13.6 
2.5.7 
34.3 

9.60 
16.0 

13.6 
4.80 
60.0 
70.2 

65. 5 

33.5 
21.6 
16.0 
12.8 
26.4 
16.5 

12.1 
25-26 
14 
10 

6-« 


69 

40.6 

18.8 

39.1 

10.1 

9.42 
15.5 

5.50 
13.0 
24.3 
36.5 

12.3 
15.2 

11.7 
7.10 
64.5 
69.6 
62.3 

25.5 
21.7 
14.8 
12.0 
24.3 
16.2 

13.0 
25-26 
11 


0-0 


40 

42.0 

22.2 

35.0 

11.5 

10.5 
12.8 
5.25 
12.7 
25.8 
38.7 

12.5 
12.3 

10.5 
9. 25 
65.0 
74.0 
65.0 

29.0 
20.0 
14.3 
13.0 
20.3 
18.3 

1.5.5 
25-25 
11 


0-7 


33 


Len^h of head 


41 2 


Greatest depth of body 

Greatest width of head 

Lenf;th of caudal podunclo 

Least depth of caudal pe- 

Hnnrln 


13.7 
36.4 
7.58 

97 


Length of snout 


11 2 


Diameter of eye 


7 68 


Width of intororbital space. .. 

Postorbital length of head 

Length of disk 


9.40 
22.8 
32 4 


Distance from base last dorsal 

ray to midcaudal fln base... 

Gape or tip of snout to rictus. . 


9.70 


Distance from rear margin of 
disk to anus 


11 2 


Anus (center) to anal origin... 

Snout tip to dorsal origin 

Snout to anal origin 


10.9 
69.1 
77 3 


Snout to anus 


63 7 


Length of dorsal fln base (to 
base of last ray) 


25 8 


Length of anal fln base 

Longest ray of dorsal fln 

Longest ray of anal fln 

Longest ray of caudal fln 

Longest ray of pectoral fln 

Length of third ray from bot- 
tom of p*«toral On 


16.7 
11.2 
11.5 
20.4 
12.4 

11.6 


Numijer of pectoral fln rays... 
Dorsal rays 


22-22 
10 


Anal rays 


8 


Number of upper pectoral 
rays above upper edge of 
attachment of gill mem- 
branes 


6-0 







See table 3 for measurements made on two specimens from Brazil. 

I have cxaiiiinod tlie type of Gohirsox ynjiui Nichols and find that it 
jjosse.sses the harbcllike .structures around the mouth and in other 
respects resembles Cotylis nigripinnis nigHpinnis to which I refer it 



68 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

as a synonym. The teeth of the lower jaw at the front have uneven 
edges but are not bifid or trifid as in certain other genera. I count 
dorsal rays as 12, anal as 9, and pectoral 23. 

The following two collections, referred to this species with uncer- 
tainty, contain very small specimens that do not show certain charac- 
ters fully developed and may represent an undescribed species of 
small size : 

U.S.N.M, No. 83862, 4 specimens, 10.5 to 11 mm., Trinidad, Albatross, January 
SO-February 21, 1884. 

U.M.M.Z. No. 131173, 4 specimens, 8.3 to 9.5 mm., Velasco, Tex., Rice Institute, 
April 17, 1923. 

Material examined. — Maeyland and Virginia: U.S.N.M. Nos. 
30400, 30407, 43064, 68829, 67760, 67761, 68391, 74852, 76530, 76531, 
77929, 83593, 85087, 85681, 86313, 88583, 88586, 89340, 91210-91239, 
92024, 92031, 93759, 93805, 104930, 109846, 122392, 122394, totaling 420 
specimens. North Carolina: U.S.N.M. Nos. 4905, 85088, 122395, 
122396, total 4 specimens. South Carolina: U.S.N.M. Nos. 26311, 
59053, 59061, totaling 9 specimens. Florida: U.S.N.M. Nos. 26611, 
30471, 30861 (2 cotypes of virgatulus) , 32760, 34719, 34725, 73250, 85089, 
85090, 91456, 92213, 92215, 93716, 93882, 94896, 116933-116935, 125493, 
totaling 31 specimens. Alabama: U.S.N.M. No. 73545, 2 specimens. 
Mississippi : U. S. N. M. No. 32625, 1 specimen. Louisiana : U.S.N.M. 
Nos. 86134, 122393, 124979, totaling 3 specimens; U.M.M.Z. No. 128860, 
3 specimens. Texas: U.S.N.M. Nos. 69347, 69348, 118542, totaling 4 
specimens; U.M.M.Z. Nos. 111746 and 114471, totaling 15 specimens. 
Brazil: U.S.N.M. Nos. 87752 and 87798, totaling 3 specimens. 

Range. — Chesapeake Bay to Brazil ; West Indies. 

COTYLIS NIGRIPINNIS WOODSI, new subspecies 

Holotype. — F.M.N.H. No. 41974, a specimen 33 mm. in standard 
length, from Cocos Island at Wafer Bay, collected February 23, 1941. 

Description of only known specimen. — Detailed measurements were 
made and these are recorded in hundredths of the standard length in 
table 3. 

Head contained about 2^/^, disk 3, greatest depth of body about 7, in 
standard length ; eye equal to bony interorbital space and 1% in fleshy 
interorbital space; disk about 1.3 in head; tip of chin to front of disk 
about % leng-th of disk ; distance from base of last dorsal ray to mid- 
base of caudal fin 2.7 in length of base of dorsal fin; dorsal origin 
equidistant between midbase of caudal fin and base of upper pectoral 
ray; tips of rays of depressed anal fin reaching a little past a line 
through base of caudal fin and depressed dorsal fin not reaching quite 
to that line; size and arrangement of papillae around mouth essentially 
as described for nigripinnis ; about three pairs of incisorlike teeth at 



REVISION OF AMERICAN CLINGFISHES — SCHULTZ 09 

front of lower jaw projecting obliquely forward, followed laterally 
by one or two somewhat enlarged conical teeth, then posteriorly by a 
short row of small conical teeth ; inside of larger outer row of teeth 
a few smaller ones at front of lower jaw ; upper jaw with conical teeth, 
those at front a little enlarged; none of the teeth with trifid tips; front 
of upper jaw inside of outer teeth with a few minute teeth ; interorbital 
space flat ; each anterior nostril with a bifid dermal flap on posterior 
margin ; shoulder girdle with a fleshy lobe on its lower margin under 
gill cover; base of pectoral fin with a fleshy lobe, the posterior and 
ventral margins with a free edge that extends to the attachment of the 
opercular membrane, both of which are fused opposite the base of the 
sixth pectoral fin ray; upper edge of axial dermal flap behind pectoral 
fin fused to base of fin opposite the sixth pectoral ray; pelvic fins at- 
tached near base of about fourth pectoral fin ray; lower rays of pec- 
toral fin nearly as long as middle rays ; margins of disk and pelvic pads 
with low flattened papillae; anus a trifle closer to anal origin than to 
rear margin of disk. 

Coloration. — General coloration pale brownish in alcohol, with five 
wide indistinct bars on body, the paler interspaces narrower than 
eye ; sides of body with several very narrow pale lines; a dark elongate 
spot behind eye and a few narrow pale lines radiating posteriorly from 
orbit across gill cover; median fins black with tips of rays white. 

Rnnarkft. — This new subspecies is the representative of a similar 
form in the Atlantic from Maryland to Brazil herein recognized under 
the n;inie 7Uf7ripinnift. From that form iroodsi may be distinguished 
by a larger eye and a less deep body, as indicated in the key. 

Named voodul in honor of Lt. Loren P. Woods, U. S. N. R., who 
tentatively suggested this specimen to be an undescribed species when 
he learned that I was studying the American clingfishes. Described 
with the permission of the authorities of the Chicago Natural History 
Museum. 

COTYLIS PINNIGER (Gilbert) 

OohtcHox pinnigcr Gn.BERT, Vroc. U. S. Nnt. Mus., vol. ];i, p. W, 1890 (Puerto Re- 
fuKld. AriKf'l Islnrid. San Luis GotizaloH I'ay, and La Paz, Gulf of California). — 
Ilxr.EfRiN, P.ull. Mns. Hist. Nat. Paris, vol. 7, ii. 1<;2. 1001 (fJiilf of r'alifornin). 

Brj/nxrlfirrf X pitinif/rr .Fobdan and Evkbmnnn, I'roc. California Acad. Sci., vol. 0, p. 
2.^ pi. .'51. ISrtf,; IN-p. t:. S. Conin*. Fish and Fisli. for m'tr>. App.. p. 401, 1806 
(Gulf of California) ; U. S. .Nat. Mns. I'.ull. 17, p. 12.12S. ISOS (Gulf of Cali- 
fornia). — JouD.VN, EvFUMANN, and CiJVKK, Hep. U. S. Conini. Fisl). for 1028, 
pt. 2, p. 4.S8, 10.30 (Gulf of California). -KnFOJiat, Bull. Binpham Oconnogr. 
Coll., vol. 2. art. 3, p. 4H, 10.36 (Puerto Krfupio ; Gonznp) Ba.v). 

ReTnarTcH. — The longer base of the dorsal fin is not considered of gen- 
eric significance in view of other related species with dorsal fins of 
nearly the same length. 



70 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

Material examined. — Gulf of California: U.S.N.M. Nos. 44377 
(type of pinniger), 46694 (4 cotypes of pinniger), 126808 (25 cotypes 
of pinniger) , totaling 30 specimens ; F.M.N.H. No. 3338, 19 specimens. 

Range. — Gulf of California. 

Genus SICYOGASTER Brisout de Barneville 

Sicyogaster Brisout de Baenevuxe, Rev. Zool. Soc. Cuv., vol. 9, p. 144, 1846. 

(Genotype: Ooiiesox marmoratus Jenyns.) 
Caularchus Giix, Proc. Acad. Nat. Sci. Philadelphia, vol. 14, p. 330, 1862. 

(Genotype: Caularchus reticulatus=^Lepadogaster rcticulatus Girard.) 

SICYOGASTER MARMORATUS (Jenyns) 

Oobiesox marmoratus Jenyns, The zoology of the voyage of H. M. S. Beagle, 
pt. 4, Fishes, p. 140, pi. 27, figs. 1, la, lb, 1842.— Gunther, Catalogue of the 
fishes in the British Museum, vol. 3, p. 504, 1861 (Chile). — ?Abbott, Proc. 
Acad. Nat. Sci. Philadelphia, 1899, p. 363 (Peru).— Delfin, Rev. Chilena 
Hist. Nat, vol. 3-4, p. 91, 1901 (Algarrobo; Chauaral ; Punta Arenas; Cal- 
buco ; Iquique; Isla de Juan Fernandez). — Tortonese, Bol. Mus. Zool. Anat. 
Comp. Univ. Torino, vol. 47, p. 206, 1939 (Valparaiso). 

Sicyogaster marmoratus Beisout de Baenevuxe, Rev. Zool. Soc. Cuv., vol. 9, p. 144, 
1846 (Chile). 

Cotylis marmoratus MvIi-lee and Troschel, Horae ichthyologicae, pt. 3, p. 19, 
1849 (Chile). 

Remarks. — The following notes on the type of Gohiesox marmora- 
tus in the British Museum from "Archipelago of Chiloe," made by Dr. 
W. H. Longlefy, were kindly turned over to me by Dr. S. F. 
Hildebrand : 

Two specimens of T. L. [total length] 56 and 64 mm. considerably macerated, 
the smaller better preserved. D. 12; A. 10; the last anal ray missing, but its 
support still evident. The pectoral both sides with 23 rays including the rudi- 
mentary one above. In the larger fish D. 12, A. 11. 

In the small fish again I found that the membranous structure at pectoral 
base is evident for only half the vertical height of the fin but that in the lower 
half, where it is present, it exists as a very evident, freely projecting lobe. 

On very careful examination, I find that the opercular cleft extends dorsally 
about to the base of the upper pectoral ray. 

Material examined. — Peru: U.S.N.M. No. 101706, 1 specimen. 
Chiles U.S.N.M. Nos. 77381, 88819-88821, 88824, totaling 6 specimens. 
Range. — Peru and Chile. 

SICYOGASTER MAEANDRICUS (Girard) 

Lepadogaster reticulatus Girard, Proc. Acad. Nat. Sci. Philadelphia, 1854, p. 

155 ( San Luis Obispo, Calif . ) (preoccupied). 
Lepadogaster maeandricus Girard, Explorations and surveys for a railroad 

route from the Mississippi River to the Pacific Ocean, vol. 10, pt. 4, p. 130, 

1858 (San Luis Obispo, S. Faralones, Calif.) (new name). 
Oobiesox maeandricus GiiNTHER, Catalogue of the fishes in the British Museum, 

vol. 3, p. 505, 1861 (Monterey). 
Caularchus reticulatus Gill, Proc. Acad. Nat. Sci. Philadelphia, vol. 14, p. 330, 

1862. 



REVISION OF AMERICAN CLINGFISHES — SCHULTZ 71 

Gobicsox rcticulatun Jordan and Joi'Y, Proc. U. S. Nat. Mus., vol. 4, p. 5, 1881 
(Monterey and Cniie Flattery). — Joiu)an and GiiJiUix, Proc. U. S. Nat. Mus., 
vol. 4. p. 63. ISSI (Monterey to Pu.uet Sound).— Rosa Smith, Proc. U. S. 
Nat. Mu.s., vol. 4. p. 140, 141. iSSl (San Diego). 

Caularchus nincaiuiricus Joiu)an and Evkkma.nn. Rep. U. S. Comm. Fish and 
Fish, for ISO'), App., p. 491, 1S96 (Vancouver Island to Monterey) ; U. S. 
Nat. Mus. Bull. 47, pt. 3, p. 2328, 189S (Vancouver Island to Point Concep- 
tion).— Ouitel, Arch. Zool. Exper., vol. 5, No. 5, pp. 625-()39, 1906 (anat- 
omy). — EvKEMANN and GoiJ)SB()KOUGH, Bull. U. S. Bur. Fish., vol. 26 p. 
336, 1907 (Fort Rupert; Gabriola Island). — Joudan, E\'ei:mann. and Clabk, 
Rep. U. S. Comm. Fish, for 1928. pt. 2, p. 4S8. 1930 (Vancouver Island to 
Monterey). — Schultz, Keys to the tishes of Washington, Oregon and closely 
adjoining regions, ed. 1, p. 197, 1930 (British Columbia to Point Arguello, 
Calif.). — ScHULTZ and DeLacey, Journ. Pan-Pacific Res. Inst. (Mid-Pacific 
Mag.), July-September, 1936, p. 211, 213 (British Columbia to Point 
Arguello, Calif.) (see this reference for additional references). — Wilby, 
Copeia, 1936, p. 116 (British Columbia). 

Material examined. — British Columbia : U.S.N.M, Nos. 49083, 
60.548, 60549, 64022, 82153, 82154, 103563, 103564, 103566, 103567, 120446, 
120447. 12G811. totalin*!; 64 specimens. Washington: U.S.N.M. Nos. 
23405, 27329, 3S334, 42049, 83208, 83964, 103565, totaling 32 specimens. 
Ori:gon: U.S.N.M. No. 91974, 1 specimen. California: U.S.N.M. 
Nos. 516 (type of reticulatus^maeandricus) , 101382, 101383, 101388, 
totaling 4 specimens. 

Range. — Queen Charlotte Islands to San Diego, Calif. ; Puget Sound. 

ARCOS, new genus 

Genotype. — Gobiesox erythrops Gilbert. 

This genus is characterized by the groove along the anterior or upper 
margin of the preniiixillary which arches in a convex manner over the 
tip of the snout; the orbits are larger than in any other genus of Amer- 
ican clingfishes, their diameter much greater than the least width of the 
bony interorbital. In addition, the axial dermal flap bcliind the pec- 
toral lin has its dorsal edge attached mucii above the niidbase of pec- 
toral ; the incisorlike teeth at front of lower jaw have smooth tips, and 
lhe.se teeth project forward horizontally and do not oppose the teeth 
at front of upper jaw, whicli are nearly conical; there are no papillae 
around the mouth, although the usual lobelike ridges occur on lower 
jaw and chin; gill membranes are joined at upper edge of pectoral fin 
ba.se or appear to be opposite base of first pecloral fin ray; the free 
posterior margin of fleshy pad on outer surface of pectoral base is con- 
fined to the lower half of that fin and not iibove the thirteenth ray from 
the top. Other characters are tlio.se of the genotype. 

Names Arcos in reference to the arched groove on the tip of the 
snout. 



72 PROCEEDINGS OF THE NATIONAL MUSEUM vol.98 

ARCOS POECILOPHTHALMUS (Jenyns) 

Gohiesox poecilophthalmus Jenyns, The zoology of the voyage of H. M. S. Beagle, 
pt. 4, Fishes, p. 141, pi. 27, fig. 2, 2a 2b, 1842 (Chatham Island).— GtJNTHER, 
Catalogue of the fishes in the British Museum, vol. 3, p. 503, 1861 (Chatham 
Island). — Jordan and Eveemann, Rep. U. S. Comm. Fish and Fish, for 1895, 
App., p. 491, 1896 (Chatham Island) ; U. S. Nat. Mus. Bull. 47, pt. 3, p. 2335, 
1898 (Chatham Island). 

Cotylis poecilophthalnius Mtjller and Teoschel, Horae ichthyologicae, pt. 3, p. 19, 
1849 (Galapagos). 

Tomicodon poecilopMMlmos Bbisout de Barne\ille, Rev. Zool. Soc. Cuv., vol. 9, 
p. 144, 1846. 

GoUesox erythrops Jordan and Gilbert, Proc. U. S. Nat. Mus., vol. 4, p. 360, 1882 
(Mazatlan). — Jordan and Evermann, Rep. U. S. Comm. Fish and Fish, for 
1895, App., p. 491, 1896 (Mazatlan ; Tres Marias Island) ; U. S. Nat. Mus. Bull. 
47, pt. 3, p. 2336, 1898 (Mazatlan) . — Jordan, Evermann, and Clark, Rep. U. S. 
Comm. Fish, for 1928, pt. 2, p. 490, 1930 (Mazatlan; Tres Marias Island). 

GoUesox paradiseus Herre, Publ. Field Mus. Nat. Hist., zool. ser., vol. 18, p. 432, 
1935 (Eden Island ; South Seymour Island) ; vol. 21, p. 393, fig. 36, 1936 (Eden 
Island; South Seymour Island). 

Remarks. — This species is recognizable by its very large eyes and 
narrow interorbital space. It is a small species and usually red in 
color. 

Dr. S. F. Hildebrand kindly turned over the following note by Dr. 
W. H. Longley on the type of Gohiesox 'poecilophthalmus from Chat- 
ham Island : 

T. L. [total length] 45 mm. D. 8, A. 7, P. 21+ rod [or 22 rays]. 

Material examined. — Mazatlan: U.S.N.M. No. 30885 (type of 
erythrops). Panama Bay (Secas Islands) : U.S.N.M. No. 101708, 5 
specimens. Galapagos Islands: U.S.N.M. No. 65516, 1 specimen; 
F.M.N.H. Nos. 17404 and 17405 (type and paratype of paradiseus) . 

Range. — Mazatlan to Panama and Galapagos Islands. 

ARCOS MACROPHTHALMUS (Giinther) 

GoMesox macropMhalmus Gunthee, Catalogue of the fishes in the British 
Museum, vol. 3, p. 502, 1861 (habitat unknown) [probably West Indies]. — 
Jordan and Evermann, U. S. Nat. Mus. Bull. 47, pt. 3, p. 2335, 1898 (St. 
Thomas). — Metzelaar, Bijd. Dierk. Feest. Num. 70th Geboortedag van Dr. Max 
Weber, pt. 22, p. 140, 1922 (Caracas Bay).— Beebe and Tee- Van, Zoologica, 
vol. 10, No. 1, p. 251, fig., 1928 (Lamentin Reef, Port-au-Prince Bay, Haiti). — 
Jordan, Evermann, and Clark, Rep. U. S. Comm. Fish, for 1928, pt. 2, p. 489, 
1930 (probably West Indies). 

Gohiesox cerasinus Cope, Trans. Amer. Philos. Soc, vol. 14, p. 413, 1871 (St. Mar- 
tins, West Indies).— Jordan and Evermann, Rep. U. S. Comm. Fish and Fish, 
for 1895, App., p. 492, 1896 (St. Martins) ; U. S. Nat. Mus. Bull. 47, pt. 3, p. 2336, 
1898 (St. Martins). — Fowler, Proc. Acad. Nat. vSci. Philadelphia, vol. 71, p. 
143, 149, 1919 (St. Martins; Jamaica).—? Metzelaae, Bijd. Dierk. Feest. 
Num. 70th Geboortedag van Dr. Max Weber, pt. 22, p. 140, 1922 (Caracas 
Bay). — Jordan, Eveemann, and Clark, Rep. U. S. Comm. Fish, for 1928, pt. 
2, p. 489, 1930 (St. Martins, West Indies). 



REVISION OF AMERICAN CLINGFISHES — SCHTJLTZ 73 

Gobicsox tudes Bvtujmann and Maksh. Bull. U. S. Fish Couun., vol. 20, pt. 1, p. 

305, 1900 (Culebru. Puerto Rico). 
Sicyancs yumurina Rivebo, rroc. Boston Soc. Nat. Illst., vol. 41, No. 4, p. 74, 

1936 (Matanza.s, reef at entrance of the Bay). 
Ouhiesox androi^icHxis Rcsen, Lunds Univ. Arf^-Skr., now .sor. (Afd. Math. Nat.), 

vol. 7. No. 5, p. 6."), 1911 (Ma.'^tic Point, Andros; Nas.sau. Bahamas). — Jokdan, 

EvEKMANN. and Clabk, Rep. U. S. Comm. Fi.<;h. for 1928, pt. 2, p. 4S9, 1930 

(Bahamas). 
Gohienox androHiensis Rosen=[not] Oobiesox riihifjinosus (I'oey), Longley, 

Carnegie Inst. Washington, Year Book, No. 34, p. 284, 1035. 
Oobicsox ccphalus Metzei^vak, Bijd. Dierk. Feest. Num. 70th Geboortedag van Dr. 

Max Weber, pt. 22, p. 139, 1922 (Caracas Bay) (see comment below). 
Gobicsox ma:'rophthaU}ius Jordan and E\'ermann, Rep. U. S. Comra. Fish and 

Fish, for 1895, App., p. 492, 1896 (West Indies). 

Re77ia7'A-s. — Tliis species has the largest eyes of any American form 
in the Athintic and is red in color when alive. The eyes are much 
wider than the narrow int^rorbital space. 

Tlirough the kindness of Dr. Thomas Barbour I have examined a 
paratype of Sicyases yumurina Kivero and refer it to this species. 
Some traces of the red color still remain on this specimen. 

Dr. S. F. Hildobrand very kindly turned over to me the following 
wuiQs, by Dr. W. H. Longley made on the type of Gohiesox niaci'oph- 
thalmus Giinther in the British Museum: 

T. L. [total length] .'4 mm. D. 8; A. 7. First ray in each tin very slightly 
tiiamentous. I'. 22 and a short, vestigial uijper 23rd. Same on both sides. The 
outline of the fin rounded. A strong subopercular spine with a deep groove on 
Its ventral surface and reaching beyond the base of any of the pectoral rays. 
The membranous sac at the ba.se of the pectoral extends upward only to the 
base of the eighth ray counting up from the ventral margin, but is a very evident 
structure. The opercular cleft is of the full width of the iMX-toral base and 
extend.s up to the level of the upper margin of the dwarf ray, which is quite a 
sizable stub one-third the length of the secctnd. 

Eye 5.0 mm.=snout; bony interorbital — 3.0 nim.=preorbital width . . . 

The dorsal origin midway between tip of caudal and posteri(»r margin of the 
I)upil. 

Di'. Uddf'brand al.M) turned over to nie the following notes made by 
Dr. Longley in the Berlin Museum, on the probable types of Gohiesox 
an//ronirnftis collected by Rosen in the Bahamas: 

Spec. 1. T. L. [total length) 2^5 mm. 1 ►. 7. A. 6, P. 24-24. the outer ray short. 
Two or rhn^ pairs of teeth above slightly flattened. Three pairs below more 
flattened, the anterior distinctly etdarged. The bnrder of the lower inci.sors only 
slightly crernilated, more nearly tnuicate fhan on No. 3. The branchial deft ex- 
tending up to the base «»f th(! upper p<'cloral ray. No groove on the siibopercidar 
spine. The nasal cirrus is a flap half flic diameter of the narlal orifice In width. 
On one side It ends In two filaments . . , 

Spec. 2. T. L. 25 nmi. I). 8, A. 7. Gill apparatus as above . . . 

Sp«'C. 3. T. L. 22 mm. D. 0. A 7, P. 22-22. Gill apparatus as above . . . 
Anterior face of lower Incisors slightly fluted, the fre<? border of the teeth almost 
bicuspid. 



74 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

Dr. Hildebrand furnished me the following notes made by Dr. 
Longley in the Museum at Amsterdam on specimens reported upon by 
Metzelaar (1922) from Curasao (Caracas Bay) : 

Oobiesox macrophthalmus . . . much fringed nasal cirri, the wide opercular 
cleft, the incomplete fold behind it with isolated lower lobe. 

T. L. [total length] 70 mm. D. 8, A. 7, P. 24-24 including stub. 

Gohiesox cephalu>s ... Is the same as last [macrophthalmus] . . . T. L. 20 
mm. D. 8, A. 7, P. 23-23, stub included . . . The gill cleft extends entire width 
of base of pectoral fin. I get no fold at all along the fin base. The anterior 
teeth of the lower jaw are enlarged, the middle much flattened and larger than 
the next pair. 

Material examined. — Bahama Islands: U.S.N.M. Nos. 38386 and 
53220, totaling 2 specimens. Jamaica : U.S.N.M. No. 78142, 1 speci- 
men. St. Thomas : U.S.N.M. Nos. 78157 and 78158, totaling 2 speci- 
mens. San Juan Island : U.S.N.M. No. 117423, 4 specimens. Virgin 
Islands : U.S.N.M. No. 117412, 3 specimens. Martinique : U.S.N.M. 
No. 117452, 2 specimens. Puerto Kico: U.S.N.M. No. 126181, 1 
specimen. 

Range. — Bahama Islands and West Indies. 

Genus GOBIESOX Lacepede 

Oohiesox Lacepede, Histoire naturelle des poissons, vol. 2, p. 595, fig., 1800. 

(Genotype: OoMesox cephalus Lacepede.) 
Megaphalus Rafinesque, Analyse de la nature, p. 86, 1815. (Genotype: Oobiesox 

cephalus.) (Substitute name for Oobiesox.) 

GOBIESOX CEPHALUS Lacepede 

Oobiesox cephalus Lacepede, Histoire naturelle des poissons, vol. 2, pp. 595, 596, 
fig., 1800 (fresh-water rivers of South America). — Brisout de Barneyille, 
Rev. Zool. Soc. Cuv., vol. 9, p. 145, 1846 (Martinique). — Gxjnthee, Catalogue 
of the fishes in the British Museum, vol. 3, pp. 499, 566, 1861 (Caribbean 
Sea; St. Domingo; West Indies). — Jordan and Eveemann, Rep. U. S. Comm. 
Fish and Fish, for 1895, App., p. 491, 1896 (West Indies) ; U. S. Nat. Mus. 
Bull. 47, pt. 8, p. 2332, 1898 (Caribbean Sea ) .— GinxEL, Arch. Zool. Exper., 
vol. 5, No. 5, pp. 640-645, 1906 (anatomy). — Blosser, Ann. Carnegie Mus., 
vol. 6, p. 300, 1909 (St. Croix).— Fowler, Proc. Acad. Nat. Sci. Philadelphia, 
vol. 71, p. 143, 1919 (St. Martins, West Indies).— Jordan, Evermann, and 
CL-^RK, Rep. U. S. Comm. Fish, for 1928, pt. 2, p. 489, 1930 (West Indies).— 
?RiVEEO, Proc. Boston Soc. Nat. Hist, vol. 41, No. 4, p. 73, 1936 (Habana). 

Oobiesox tudes Richardson, Zoology of the voyage of H. M. S. Sulphur, Ichthyol- 
ogy, vol. 1, p. 103, pi. 46, figs. 1-3, 1844 (locality unknown ) .—Jordan and 
Eveemann, U. S. Nat. Mus. Bull. 47, pt. 3, p. 2333, 1898 (West Indies).— 
Joedan, Eveemann, and Clabk, Rep. U. S. Comm. Fish, for 1928, pt. 2, 
p. 489, 1930 (probably West Indies). 

Oobiesox tudes 'Richardson= Oobiesox cephalus Lacepfede, Longlet, Carnegie 
Inst. Washington Year Book, No. 34, p. 284, 1935. 

Cotylis stannii MtJiXER and Tboschel, Horae ichthyologicae, pt. 3, p. 18, pi. 3, 
fig. 3, 1849 (Brazil). 

Oobiesox costaricensis Meek, Publ. Field Mus. Nat. Hist., zool. ser., vol. 10, No. 
7, p. 74, 1912 (Zent River, Atlantic drainage, Costa Rica). — Behre, Ann. 



REVISION OF AMERICAN CLINGFISHES — SCHULTZ 75 

Carnegie Mus., vol. IS, p. 314, 192S (tributary to Rio Cricatuola near Con- 
quautu). — JOKOAN, Evkkmann, and Ci-\kk, Kep. U. S. Couim. Fish, for 1U2S, 
pt. 2, p. 4S9, 1930 (Costa Kica). 
Gobiesox ramsdeni RnxEo, Proc. Boston Soc. Nat. Hist., vol. 41, No. 4, p. 73, 
193G (Rio Toa. "El Palenque" Yateras, GuautAnanio, in Orionte Province, 
Cuba ) . 

Remarks. — Dr. S. F. Hildebrand kindly turned over to me the fol- 
lowintr note made by Dr. W. H. Longley in the Paris Museum of 
Natural History on a specimen, No. 513-1:, of Gobiesox cephalus Lace- 
pede: 

T. L. [total length] 9.") mm., D. S, A. G, P. 21 and a stub on outer sitle. If any 
are Lacei)ttde's specimens, this must be it, others are all too late. 

Contrary to most references in the literature, Lacepede did not report 
G. ccphalus from the ''Caribbean Sea" but from fresh-water rivers of 
South America. Since Lacepede mentions fresh water once and rivers 
twice in his dc-^cription as the habitat of this species, I assume he did 
not make a mistake in the locality where the species occurred. 

1 have before me a series of specimens of Gobiesox from fresh-water 
streams of northern South America and Central America on the At- 
lantic side, and these agree in most characteristics throughout the area 
represented. Noteworthy is the arrangement of the teeth. On the 
lower jaw anteriorly the teeth of the outer row are somewhat enlarged, 
short, narrow and incisorlike, not crowded or projecting forward at the 
symphysis; front of upper jaw with conical teeth; lateral teeth in 
both jaws conical ; sometimes with one or two enlarged caninelike teeth 
at front sides of lower or upper jaws; upper jaw with an inner patch 
of small conical teeth ; origin of dorsal fin usually equidistant between 
midbase of caudal fin and tips of pectoral fin rays; anus usually equi- 
distant between anal origin and rear margin of disk or a little closer to 
anal origin ; anal origin behind middle of bases of dorsal fin rays or 
under the liftli or sixth ; anus slightly in front of a vertical line through 
dorsal origin; head 2.2 to 2.G; disk 2.G to 3.2; depth 4 to 5.5, all in 
standard length; interorbital equals snout. 

There is a black blotch near base of dorsal fin on first rays that ap- 
pears to occur constantly on the specimens exiunined. 

William C. Schroeder, Mu.seum of Comparative Zoology, kindly 
checked the type of Gobiesox raiiisdeni (M. C. Z. No. 34152) and made 
the following observations: 

Posterolateral teeth c>f lower jaw more rnniiiclike and not smaller than front 
tf^eth; no papilla on upiH'r or lower jaws (unless I overlooki-d this) ; length of 
disk e^juals disk to midbase of anal; anus closer to anal origin than to rear 
ninrpln of disk by an i'ye's diameter. Standard length 107 [mm.]; head 43; 
length of disk 30; depth of body 25; D. 8; A. 5 or 6; I'. 20; eye .^; interorbital 10. 

The foregoing counts may be considered as correcting those given in 
the original description of ramsdeni. 



76 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

Matenal examined. — ^West Indies: U.S.N.M. Nos. 26479, 29848, 
93793, totaling 11 specimens. Costa Rica : U.S.N.M. No. 74246 (para- 
type of costarwensis) ; F. M. N. H. Nos. 7677 and 7813 (type and 2 
paratypes of costaricerisis) , 7814, 7815, totaling 6 specimens. Vene- 
zuela (Macuto): U.S.N.M. Nos. 93815-93817, 93820-93822, 93827, 
totaling 16 specimens. Colombia (Eio Dagua) : U.S.N.M. No. 120217, 
1 specimen. 

Range. — ^West Indies; Costa Rica to Brazil. 

GOBIESOX FULVUS Meek 

Gobiesox fulvus Meek, Publ. Field Columbian Mus., zool. ser., vol. 7, No. 5, p. 149, 
1907 (Cocos Island). 

Remarks. — The following specimens were studied: Cocos Island: 
F. M. N. H. No. 6035 (type of fulvus) ; U.S.N.M. No. 91832, 1 speci- 
men ; U. M. M. Z. Nos. 131512 and 131513, 2 specimens. 

Range. — Cocos Island. 

GOBIESOX PUNCTULATUS (Poey) 

Sicyases punctulatus Poet, Enumeratio piscium Cubensium, p. 124, 1876 
(Cuba). — Jordan, Evekmann, and Clark, Rep. U. S. Comm. Fish, for 1928, pt. 
2, p. 490, 1930 (Cuba). 

Gobiesox punctulatus Jordan and E\'ekmann, Rep. U. S. Comm. Fish and Fish, for 
1895, App., p. 492, 1896 (Cuba) ; U. S. Nat. Mus. Bull. 47, pt. 3, p. 2338, 1898 
(Cuba). — Metzeiaak, Bijd. Dierk. Feest.-Num. 70th Geboortedag van Dr. 
Max Weber, pt. 22, p. 140, 1922 (Caracas Bay). 

Gobiesox Jiaeres Jordan and Bollman, Proc. U. S. Nat. Mus., vol. 11, p. 552, 1889 
(Green Turtle Cay, Bahamas). — Jordan and Evebmann, Rep. U. S. Comm. 
Fish and Fish, for 1895, App., p. 492, 1896 (Green Turtle Cay, Bahamas) ; 
U. S. Nat. Mus. Bull. 47, pt. 3, p. 2337, 1898 (Green Turtle Cay, Bahamas). 

Gobiesox nudiis Metzelaar, Report on the fishes collected by Dr. J. Boeke in the 
Dutch West Indies 1904-1905, pt. 1, p. 151, 1919 (Curasao and Bonaire). 

Gobiesox l[h]aeres Jordan and Bollman=Gf. punctulatus (Poey), Longley, Car- 
negie Inst. Washington, Year Book, No. 33, p. 271, 1934. 

Gobiesox vittatus Mepzexaae, Bijd. Dierk. Feest.-Num. 70th Geboortedag van 
Dr. Max Weber, pt. 22, p. 140, fig. 3, 1922 (Caracas Bay ).— Jordan, Evekmann, 
and Clark, Rept. U. S. Comm. Fish, for 1928, pt. 2, p. 489, 1930 (Curagao). 

Sicyases haeres Jordan, Eveemann, and Clark, Rep. U. S. Comm. Fish, for 1928, 
pt. 2, p. 490', 1930 (Green Turtle Cay, Bahamas). 

Remarks. — This is a small species and may be recognized by its 
relatively short, thick and deep body giving it the appearance of 
robustness. 

Dr. S. F. Hildebrand kindly turned over to me the notes made by 
Dr. W. H. Longley in the Museum at Amsterdam on the type of 
Gobiesox vittatus Metzelaar, which follow : 

T. L. [total length] 29.0 mm., D. 11, A. 7. P. 1^20, orbit 2.0 mm. Interorbital 
2.0 mm., anterior margin of sucking disk crenulated, the units not stalked. The 
nasal cirrus small, expanded, doubly pointed. Opercular cleft extending up to 



REVISION OF AMERICAN CLINGFISHES — SCHULTZ 77 

base of 5th pectoral ray, tin' fold behind it complete, with a distinct lobe below. 
The origin of the dorsal midway between tip of snout and of tip of caudal. The 
anterior teeth above simple, sharp-pointed, circular in cross-section, I think. 
Three pairs anterior teeth below flattened, not truncate, but leaf-shaped or even 
a little more spatulate. 

Body pretty uniformly covered with dark chromatophore.s at an average distance 
from one another of twice their diameter . . . 

Material examined. — Bahama Islands: U.S.N.M. No. 41733 (type 
of hacres). Cuba : U.S.N.M. No. 37.->31, 5 specimens. Tkxas : U.S.N.M. 
No, 1219G2, 1 specimen. British Honduras: U.S.N.M. No. 91816, 
1 specimen. Locality unknown: U.S.N.M. No. 34442, 1 specimen. 

Range. — Bahama Islands and West Indies; Texas and British 
Honduras. 

GOBIESOX ADUSTUS Jordan and Gilbert 

GobiesoT adustus Jobi>.\n and Gilbert, Proc. U. S. Nat. Mus., vol. 4, p. 360, 1882 
(Mazatlfin) ; vol. .">, p. 627, 1883 (Central America). — .Joudan and Evebmann, 
Kep. U. S. Comm. Fish and t^sh. for 1805, App., p. 491, 1896 (Mazatian); 
U. S. Nat. Mus. Bull. 47, pt. 3, p. 2334, 1898 (Mazatlfin; Pacific coa.st of 
Mexico. — JoKUAN, Evermann, and Clakk, Rep. U. S. Comm. Fish, for 1928, 
pt. 2, p. 489, 1930 (Pacific coast of Mexico). 

Material examined. — U.S.N.M. No. 29249 (3 types of adustus).^ 
Mazatlan, C. H. Gilbert. 
Range. — Mazatlan, Mexico. 

GOBIESOX FUNEBRIS Gilbert 

Gohuxox lunehrxH Giijiert, Proc. U. S. Nat. Mu.s., vol. 13, p. 95, 1890 (Puerto 
Kefugio, Angel Island, and La Paz, Gulf of California). — Jordan and 
E\xbmann, Rep. U. S. Comm. Fish and Fish, for 1895, App., p. 491, 1896 
(Gulf of California) ; U. S. Nat. Mus. Bull. 47, pt. 3, p. 2334, 1898 (Angel 
Island and La Paz, Gulf of California). — Jordan. Evermann, and Clark, 
Rep. U. S, Comm. FisJi. for 1928, pt. 2, p. 4.S9, 1930 (Gulf of California).— 
Bredrr, Bull. BiiiKham Oceanogr. Coll., vol. 2, art. 3. p. 48, 1936 (Puerto 
Refupio) . 

Rem/irkf(. — This species may be recofjnized by its small disk, the at- 
tachment of the gill membranes opposite the bases of the fifth or 
sixth pectoral fin rays, and the poor development of the fleshy pad on 
otitor pectoral fin base, along with its narrower head. 

Material examinrd. — (julf of California: U.S.N.M. Nos. 44378 
(type of funebris). 119720, 124955 (2 cotypes of funehris), totaling 4 
specimens; F.M.N.H. No. 8997, 2 specimens. 

Range. — Gulf of California. 



u I eovmNiii NT miNTiNs orricti t*4« 



PROCEEDINGS OF THE UNITED STATES NATIONAL MUESUM 



issued hlik^ V'gl^l hy the 




SMITHSONIAN INSTITUTION 
U. S. NATIONAL MUSEUM 

Vol. % Waihington: 1945 No. 3188 



NEW BEETLES OF THE FAMILY EUCNEMIDIDAE FROM 
CENTllAL AMERICA AND THE WEST INDIES 



By W. S. Fisher 



During the process of rearranging the American species of the 
family Eucnemididao (Coleoptera) in the United States National 
Mu>eum, a number of new genera and species were found. These are 
described herein. According to Fleutiaux (Rev. Franc. Ent., voL 2, 
p. 1, 1935), the name Eucnemididae sliould be used for this family in- 
stead of Melasidae. 

Genus TEMMLLUS Bonvouloir 

TEMNILLUS ASITRICOLUS, new species 

Oval-oblong, dark reddish brown, the antenna sliglitly paler, opaque, 
glabrous. 

Head in-egiilarly convex, with an irregular, deep, transverse depres- 
sion on vertex and a very deep, irregular depression between antennae, 
coarsely, deej)ly, densely punctate, the intervals vaguely granulose; 
clypeus short, wide, anterior margin strongly, obtusely toothed at 
middle, broadly, arciiately cniarginaU' on each side. Antenna not 
exieiidiiig to base of j)roiiotum, comjiact ; first segment large, irregular, 
slightly flattened, as long as following five segments united, with a 
sharp tooth on underside at apex; segments -2 to 10 wider than long, 
each with two round (lej)ressions on npi)er surface; segment 11 
obliquely truncate at apex. 

PionotMHi distinctly widei- than long, strongly conv«'.\, with a inir- 
Tow, longit ndinal, median gi-oove extending fi'oin base to apex and a 
small, deeper depression at middle of groove; sides nearly parallel, 
slighth' sinuate along basal two-thirds, bioadly ronnded anterinily ; 
base truncate, posterior angles strongly piojecting backward ; surface 
r.'jdiijii tr. ~,j 



80 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

coarsely, asperately punctate at middle, ocellate-punctate toward sides. 
Scutellum square, coarsely j)unctate at middle, broadly rounded at 
apex. 

Elytra very strongly convex, strongly striate, the striae deeper near 
apex ; sides parallel from bases to apical third, then arcuately narrowed 
to apices, which are conjointly obtusely angulated; surface densely, 
finely rugose and finely punctate, the striae coarsely, deeply punctate. 

Abdomen beneath coarsely, densely, uniformly punctate, last ster- 
nite with two round, deep depressions; prosternal process narrow, 
coarsely punctate, rounded at apex, sides nearly parallel; posterior 
coxae triangular. 

Length 9.5 mm., width 3.25 mm. 

Ty])e locality. — Verdant Vale, Arima, Trinidad, British West In- 
dies. 

T'y^e.—U.S.N.M. No. 57163. 

Remarks. — Described from a single specimen collected at the type 
locality. 

This species resembles Temnillus Uprieuri (Guerin) but differs 
from that species in being more elongate and in having the head very 
deeply depressed between the antennae and on the vertex, with the 
surface more finely punctured, the anterior margin of the clypeus 
strongly toothed at the middle, the pronotum longitudinally grooved 
with a distinct depression at the middle, and the disk asperately punc- 
tured, the elytra strongly striate with the intervals finely rugose and 
the scutellum and prosternal process coarsely punctured. 

NEODIAPODIUS, new genus 

Body elongate, subcylindrical, slightly attenuate posteriorly. Head 
large, moderately convex, boardly, transversely flattened behind clyp- 
eus; labrum invisible; eyes partially covered by prothorax; clypeus 
moderately contracted at base. Antenna short, extending slightly 
beyond base of pronotum, slightly serrate in male, strongly serrate 
in female ; second segment very small ; third segment slightly shorter 
than first ; segments 4 to 10 slightly shorter than third and subequal in 
length to one another; segment 11 narrowly oblong, acute at apex. 
Palpi abnormal, last segment of labial and maxillary palpi in male 
broadly rounded at base, nearly four times as long as wide at base, 
one-third as wide at apex as at base, with the outer margin slightly 
concave and inner margin slightly convex, in the female short, broadly 
oblong, twice as long as wide at middle, with outer margin obliquely 
trmicate on apical half and inner margin strongly convex, the outer 
surface with a round depression at middle. Pronotum as long as wide, 
rounded and slightly sinuate in front, but not carinate ; lateral mar- 
gins distinct, entire; posterior angles strongly projecting backward. 



NEW EUCNEMIDID BEETLES — FISHER 81 

Sciitelluni elongute-tritinguliir, gibbose autorioily, .sli<i:hlly emarginute 
at apex. Elytra strongly convex, attenuate posteriorly, distinctly stri- 
ate. Propleural triangle as long as ^vide at base; antennal groove 
along lateral margin shallow, not very wide, smooth along outer mar- 
gin, not margined internally. Marginal carina and prosternal suture 
converging anteriorly, the latter closed, carinate and strongly ele- 
vated anteriorly. Metasternal einmeron visible. Metasternal epi- 
tternum broad, sides parallel. Posterior coxae broad, sides parallel. 
Abdomen convex; last sternite slightly gibbose toward apex, which 
is produced into a short, broad, truncate projection. Femora com- 
pressed. Tibiae subcylindrical. Tarsi shorter than tibiae, without 
lamellae; first segment slightly shorter than following segments 
united; fourth segment hollowed out on dorsal surface for receiving 
following segment, truncate at apex; tarsal claws robust, cleft near 
apices. 

Genotype. — Neodiapodms buscki new species. 

This genus is allied to Diapodius Bonvouloir, but it differs from the 
description given for that genus in having the pronotum not longer 
than wide, the propleural triangle as long as wide at the bottom, the 
metasternal epimeron visible, the clypeus sinuate in front, the meta- 
sternal episternum broad with the sides parallel, the posterior tarsi 
shorter tlian the tibiae, the first segment of the posterior tarsus slightly 
shorter than the following segments united, the tarsal claws cleft near 
the tips, and the male antenna only slightly serrate. 

NEODIAPODIUS BUSCKI. new species 

MaJr. — Elongate, strongly convex, uniformly reddish brown, with 
the legs and palpi slightly paler, subopaque. rather densely clothed 
with very short, recumbent, whitish hairs. 

Head coarsely, confluently, rugosely punctate, without a longitudinal 
■ iirina; interocuiar carinae slightly elevated, not extending beyond 
antonnal sockets; clypeus at base twice as wide as distance between it 
and eye, in front broadly rounded at middle and sinuate on each side, 
surface longitudinally carinate at middle. 

Pronotum stroi^gly convex anteriorly, depressed along base, witli a 
more or les.s distinct longitudinal, median depression extending from 
base to near apex; sides parallel along basal two-thirds, arcuatcly 
narrowed near aj>^x; surface coarsely, confluently. rugoselv punctate. 

Elytra with sides parallel to middle, then gradually narrowed to the 
lips, which are slightly separated and acute; surface rather deei)ly. 
longitudinally striate; intervals slightly convex, densely, finely granu- 
losa' basally, becoming finely, sparsely punctate toward apices. 

Body beneath densely punctate on abdomen, more coarsely and 
deeply on i)rostei nuni, mesosternum, and melasternum; last abdominal 
sternite coarsely granulosc toward apex. 



82 PROCEEDINGS OF THE NATIONAL MUSEUM vol. se 

Length 14 mm., width 4.5 mm. 

Female.— T>\S.Qvs from the male in being larger, and in having the 
abdomen more densely clothed at the middle with more conspicuous, 
longer, erect hairs, and the last sternite more obliquely narrowed toward 
the apex, the tarsal claws cleft farther from the tips with the inner 
tooth of each claw much shorter than the outer one, and the palpi and 
antennae of different shapes. 

Length 17-20 mm., width 5.5-6 nun. 

Type locality --Lia, Chorrera, Panama. 

Type, allotype, and paratypes. — U.S.N.M. No. 57164. 

Remarks. — Described from five specimens (1 male type) collected 
at the type locality by August Busck, the male type on May 14, 1912, 
and four females on May 17, 1912. 

Genus DROMAEOLUS Kiesenwetter 

DROMAEOLUS PULCHER, new species 

Oblong, slightly convex above, moderately shining, dark brown, 
elytra and abdomen more reddish, legs yellowish brown, the elytra 
ornamented with whitish pubescent designs. 

Head slightly convex, vaguely, transversely depressed in front of 
clypeus, with a vague, short, longitudinal, smooth, median carina, 
finely, confluently punctate, sparsely clothed with moderately long, 
semierect, whitish hairs; interocular carina very strongly elevated, 
not interrupted and broadly rounded at middle ; clypeus very narrow 
at base, one-half as wide as distance to eye, broadly sinuate at apex, 
broadly depressed near anterior margin and with a strongly elevated, 
short, longitudinal carina at base. Antenna rather robust, densely 
pubescent, longitudinally carinate, not distinctly serrate; third seg- 
ment vaguely longer than second; segments 3 to 10 as wide as long 
and subequal in length to one another; segment 11 oblong, acute 
at apex. 

Pronotum slightly shorter at middle than wide at base, strongly 
convex, deflexed along base; sides arcuately converging from bases 
to apices; posterior angles strongly projecting backward along sides 
of elytra; surface densely, finely, uniformly punctate, densely 
clothed along sides and base with long, recumbent, whitish hairs, 
hairs on median part not conspicuous. 

Elytra slightly convex ; sides converging from bases to tips, which 
are conjointly broadly rounded; surface not striate, densely, finely 
punctate, densely clothed with long, semierect, blackish hairs, and 
each elytron ornamented with long, recumbent, whitish hairs as fol- 
lows : A narrow fascia along base broadly expanded at humeral angle, 
a narrow band along sutural margin connected to a broad fascia at 
middle, and a broad fascia at apex. 



NEW EUCNEMIDID BEETLES — FISHER 83 

Body beneath finely, densely punctate, rather densely clotlied %vith 
moderately long, recuini)ent, whitish haii-s; antennal grooves mar- 
ginal, narrow, equal in width for their entire length; prosternal su- 
ture not deeply grooved ; posterior coxae strongly expanded internally. 

Length 7.5 mm., width li.o mm. 

Type local it y. — Portobelo, Panama. 

fypc.—V.SS.^l. No. :.71G5. 

ReiJiarkM. — Described from a single specimen reared from a pupa 
collected in the bark of an unrecognized tree at the type locality, 
^rarch 1911, by E. A. Schwarz. 

This species resembles Dromaeolus ornatidu-s Horn but differs from 
that species in being larger, in having the elytra and abdomen reddish 
brown and the clypeus carinate at the base, and in not having the white 
j)ubescence extending along the lateral margins of the elytra or the 
elytra striate. 

DROMAEOLUS PANAMENSIS, new species 

Oblong, moderately convex, slightly shining, dark brown, the legs 
and antenna yellowish brown, rather densely, uniformly clothed 
above with short, semierect, brownish hairs, and beneath with short, 
recumbent, yellowish hairs. 

Head nearly flat, densely, coarsely ocellate-punctate, slightly' de- 
pressed near interocular carina, which is strongly elevated, arcuate, 
and not interrupted at middle; clypeus very narrow at base, one- 
third as wide as distance to eye, not carinate, broadly rounded at 
apex, the surface coarsely, confluently ocellate-punctate. Antenna 
rather robust, densely pubescent, vaguely, longitudinally carinate] 
compact, not serrate; third segment slightly longer than second; seg- 
ments 3 to 10 slightly longer than wide and subequal in length to 
one another; segment 11 oblong, subacute at apex. 

Pronotum distinctly wider than long, strongly convex, deflexed 
along ba.se, without distinct depressions or carinae; sides arcuately 
converging from ))ases to apices; surface; densely, coarsely ocellate- 
punctate. 

Klytra moderately convex; sides converging from bases to tips, 
which are conjointly broadly rounded; surface finely sti'iate, intervals 
flat, densely, finely rugose basally, finely, sparsely, punctate toward 
apices. 

I'ody beneath finely, densely punctate on abdomen, more coarsely 
ocellate-punctate on j)rosternum, mesostennnn. and metasternum ; 
nntennal grooves marginal, narrow, (h\\\'a\ in width for their entire 
length; j)rosternal suture nr)t deeply grooved; pi-oj)leural triangle 
with iK>sterior margin shorter than inner nnirgin; posterior coxae 
.strongly expanded internally. 

Length 3.4 mm., width 1.3 mm. 

Tyi>r locality. — Cabima, Pnriania. 



84 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

Type.—U.S.^M. No. 57166. 

Remarks. — Described from a single specimen collected at the type 
locality on May 28, 1911, by August Busck. 

This species is allied to Droinaeoliu moerens Horn but differs from 
the description given for that species in being dark brown, with the 
legs and the antennae yellowish brown, and in having the third seg- 
ment of the antenna subequal in length to the fourth, the elytra rugose 
at the bases, and the pronotum distinctly wider than long. 

Genus FORNAX Laporte 

FORNAX POEYI, new species 

Elongate, subcylindrical, strongly convex, slightly narrowed 
posteriorly, moderately shining, reddish brown, anterior margin of 
pronotum more reddish, and legs and antennae brownish yellow, 
rather densely, uniformly clothed with short, recumbent, yellowish 
hairs. 

Head strongly convex, without depressions or carina, coarsely, 
confluently ocellate-punctate, and more or less granulose ; interocular 
carina strongly elevated, not extending on base of clypeus; clypeus 
at base slightly wider than distance to eye. Antenna slender, ex- 
tending slightly beyond base of elytra, not distinctly serrate ; segment 
2 subequal in length to segment 4; segments 4 and 5 united longer 
than 6; the following segments subequal in length to one another; 
segment 11 oblong narrowly rounded at apex. 

Pronotum slightly wider than long ; sides nearly parallel from base 
to apex; posterior angles acute; disk moderately convex, without 
depressions or median carina ; surface rugose and finely, confluently 
ocellate-punctate. 

Elytra moderately convex; sides parallel to behind middle, then 
gradually, arcuately narrowed to tips, which are conjointly narrowly 
rounded; surface finely striate, finely, densely granulose basally, be- 
coming more sparsely, finely punctate toward apices; epipleura 
punctate, not grooved. 

Body beneath rather densely punctate, finely on abdomen, coarsely 
on prosternum, and rugosely on metasternum ; last abdominal sternite 
coarsely granulose toward apex; antennal groove narrowed by eye, 
deep, not half as wide as propleural triangle, not wider in front, the 
inner margin not very sharply defined; prosternal process flat, sides 
nearly parallel to behind coxae, acute at apex; first segment of 
posterior tarsus subequal in length to the following segments united. 

Length 4.5 mm., width 1.25 mm, 

T]/pe locality. — Cayamas, Cuba. 

Type.— U.S.^.M. No. 57167. 



NEW EUCiSTEMIDID BEETLES — FISHER 85 

Rt7narks. — Described from u sin<»:le speciiutMi collected at tlie type 
locality, January 24, by E. A. Schwarz. 

This species is allied to Fornax hdd'nis (Melsheimer) l)ut (Hirers 
from that species in bein«j: smaller and in havin<ij the head and pronotum 
finely, continently punctured, and the inner mar*rin of the antennal 
suture on the prosternum not very sharply defined. 

FORNAX VALFRIO, new species 

Oblong, subcylindrical. moderately convex, slightly narrowed pos- 
teriorly, uniformly dark, reddish brown, the legs and antennae slightly 
paler, densely clothed above with moderat^ely long, recumbent and 
semierect, brownish-yellow pubescence, which does not conceal the 
surface, beneath with shorter and less conspicuous pubescence. 

Head convex, without depressions or longitudinal carina, finely, con- 
Ihienlly punctate, finely rugose; interocular carinae slightly elevated, 
not extending on base of clypeus, the latter at base slightly wider than 
distance to eye. Antenna slender, extending to posterior coxa, not 
serrate; segments 4 to 10 similar, becoming gradually longer toward 
t ip of antenna ; segment '2 very small, shorter than 4; segment 3 longer 
than 4. 

Pronotum wider than long; sides parallel posteriorly, arcuately 
narrowed anteriorly; posterior angles subacute; disk strongly convex 
anteriorly, transversely, obliquely depressed behind middle, with a 
short, smooth line in front of scutellum, and a distinct pitlike depres- 
sion along base on each side at outer angle of scutellum ; surface finely, 
densely granulose. finely, obsoletely I'ugose anteriorly and towaid 
Literal margins. 

Elytra moderately convex; sides ])arallel to apical third, then 
slightly narrowed to the tips, which are conjointly broadly rounded, 
distinctly striate; intervals flat, finely, densnly granulose basally, be- 
coming punctate po.steriorly ; epipleurae smooth, grooved their entire 
length. 

Body beneath finely, densely punctate, more coarsely on prosterniini : 
antennal groove narrow'ed by the eye, deep, not half as wide as pro- 
I)leural triangle, not wider in front, innei- margin sliar|)ly <lelined; 
j»ro>t('rnai pi-ocess tleflexed and abruptly narrowed i)ehind anterior 
coxae, acute at apex; last abdominal sternite longilu'linally depressed 
along sides, longitudinally conipressetl ])ostei'iorIy, scabrous and acute 
at apex: first segment of posterior tarsus >Mbe(iual in length to the 
following .segments united. 

Length Il-IT) mm., width .'{.r>-4.r) mm. 

Tijpr locdlitu. — Chitaria, Cost a Ricii. 

Type and paratijpcs.—\] .f^.^ M. No. .57108. 

Rrrn/irl's. — Described fi-om thi-cc sp(>cime)is (1 typ(>) collected nt 
the type locality. December 4, HOO, by M. Valerio. 



86 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

This species resembles Fornax mendax Bonvouloir and Fornax 
ohi'utus Guerin, but it differs from both these species in having the 
pronotum transversely, obliquely depressed behind the middle, with a 
distinct pitlike depression along base on each side at outer angle of the 
scutellum, and with the surface finely granulose, and the elytra finely 
granulose basally, and it also differs from ohrutus Guerin in having the 
prosternal process deflexed behind the anterior coxae, and abruptly 
narrower to the tip, which is acute. 

Genus PLESIOFORNAX Bonvouloir 

PLESIOFORNAX NIGRINUS, new species 

Narrowly elongate, subcylindrical, strongly shining, black, with a 
faint bluish or greenish tinge, and legs brownish black, sparsely 
clothed with short, recumbent, inconspicuous hairs. 

Head strongly convex, without distinct depressions or carina, rather 
coarsely, densely punctate ; interocular carina not elevated or extend- 
ing along base of clypeus, which is strongly contracted at base, its 
width there being less than half the distance to eye, and broadly, sin- 
uately rounded in front. Antenna slender, more robust toward tip, 
extending to basal fourth of elytra, basal segments slightly longitu- 
dinally carinate; segment 2 as long as segment 5; segment 3 much 
longer than either 2 or 4 ; segment 4 longer than wide, more than half 
as long as 5, the following segments becoming gradually longer. 

Pronotum as long as wide, strongly convex; sides parallel along 
basal two-thirds, feebly, arcuately narrowed along apical third ; sur- 
face very sparsely, finely punctate, with a median, longitudinal de- 
pression extending from base to middle of pronotum, the depression 
becoming broader and deeper posteriorly. 

Elytra strongly convex; sides parallel from base to apical third, 
then arcuately narrowed to tips, which are conjointly broadly 
rounded ; surface rather coarsely, not very densely, irregularly punc- 
tate, not striate. 

Abdomen finely, densely punctate, more confluently toward apex 
of last sternite, which is obtusely angulated. Presternum sparsely 
punctate on median part, densely on propleural triangle, which is 
distinctly longer than wide at base ; antennal groove deep, smooth, and 
shining in its entire length; prosternal process broad, flat, without 
depressions, sides obliquely converging to apex, which is broadly 
rounded. Posterior tarsus with first segment as long as following 
segments united ; tarsal claws long, slender, slightly swollen near base, 
but not toothed. 

Length 11-13 mm., width 2.5-2.8 mm. 

Type locality. — Rio Hondo, Costa Rica. 

Type and paratypes.—V.S.^M. No. 57169. 



NEW EUCNEMIDID BEETLES — FISHER 87 

Remarks. — Described from three specimens (1 type). Type and 
one paratype collected at the type locality by J. Pittier, and one para- 
type collected at San Jose, Costa Rica, 1,000 to 1,200 meters, January 
15, 1934, by Ferdinand Nevermann. 

This species is allied to Ple^iofornnx loTigicorms Horn but differs 
from the description given for that species in being shining black with 
a bluish or greenish tinge and in having the antenna extending only 
(o the basal fourth of the elytra, the pronotum as long as wide, with 
the sides parallel along the basal two-thirds, the proi)leural triangle 
densely punctate, and the first segment of the posterior tarsus as long 
as the following segments united. 

Genus FARSUS Jacquelin Du Val 

FARSUS CONVEXUS, new species 

Cylindrical, robust, strongly convex, scarcely narrowed posteriorly, 
subopaqup. uniformly reddish brown, sparsely, uniformly clothed witli 
-hort, semierect, yellowish hairs, which are more recumbent on under- 
side of body. 

Head moderately convex, vaguely depressed behind clypeus, coarsely, 
confluently ocellate-punctate ; interocular carina vaguely elevated', not 
extending along base of clypeus; clypeus strongly contracted at base, 
which is nearly as wide as distance to eye, broadly sinuate or rounded 
in front. Antenna rather robust, extending slightly beyond base of 
elytra; segment 2 as long as 4, which is one-half as long as 3, the fol- 
lowing segments subequal in length to one another except 11, which is 
longer and oblong; segments 4 to 10 compact, wider than long. 

Pronotum sliglitly wider than long, strongly convex anteriorly, ob- 
liquely deflexed along base; sides parallel to near apex, then arcuately 
rounded ; anterior margin broadly rounded, distinctly carinate, the 
cai'ina curving backward on each side near apical angles; base broadly 
emarginate in fiont of scutellum; surface coarsely, densely ocellate- 
punctate at middle, scabrous toward sides. 

Elytra strongly convex, vaguely striate, the intervals rather densely, 
irregularly punctate; sides parallel to near apices, which are conjointly 
broadly rounded. 

Body beneath densely, coarsely ocellate-punctate; abdomen strongly 
convex, last sternite broadly rounded at apex. Prosterninn with 
lateral margin strongly elevated posteriorly, obsolete toward apex; 
supplementary carina sinuate, strongly elevated anteriorly, extending 
along j)rosternal suture to middle of proplenral triangle. Femora ro- 
i;usf. P<;sterior coxae strongly, angularly expanded internally. 

Length 4-6 mm., width 1.25-2 mm. 

Type lorol'ity. — Tabernilla, Caiutl Zone. 

Type and jmratyprn. — IJ.S.N.M. No. 57170. 



88 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

Remarhs. — Described from four specimens (1 type). The type and 
two paratypes collected at the type locality, May 14-17, 1907, by August 
Busck, and one paratype taken from an arc-light globe at Ancon, 
Canal Zone, during April 1911. Under this species is also included a 
specimen collected at General Ballivian, Salta Province, Argentina, 
during 1927-28, by G. L. Harrington. This specimen differs from the 
type only in having the supplementary carina on the prosternum 
vaguely indicated posteriorly and joined to the lateral carina near the 
base. 

This species is allied to Farsus oblitus Horn but differs from that 
species in being more robust, more convex, and scarcely narrowed 
posteriorly and in having the antenna shorter, with the segment 2 as 
long as segment 4, and the following segments wider than long, the 
pronotum coarsely ocellate-punctate at the middle, the sides of the 
elytra parallel to near the apices, with the surface irregularly punc- 
tate, the femora robust, the posterior coxae strongly, angularly ex- 
panded internally, and the supplementary carina on the prosternum 
strongly elevated anteriorly and extending only to the middle of the 
propleural triangle. 

^ Genus ARRHIPIS Dejean 

ARRHIPIS CUBANUS, new species 

Elongate, subcylindrical, moderately convex, rather strongly shin- 
ing, reddish brown, the antenna and legs yellowish brown, rather 
densely, uniformly clothed with short, semierect, whitish hairs. 

Head slightly convex, without depressions or carina, coarsely, 
deeply, confluently punctate; interocular carina slightly elevated, 
extending around inner margin of antennal fossa, but not along 
base of clypeus ; clypeus short, at base twice as wide as distance to eye, 
feebly, broadly rounded in front. Antenna moniliform, slightly thick- 
ened toward apex, extending to base of pronotum ; segment 3 as long 
as following two segments united ; segments 4 to 10 as wide as long, 
subequal in length to one another; segment 11 oblong, feebly incised 
on inner margin, narrowly rounded at apex. 

Pronotum as long as wide, widest near apex ; sides nearly parallel ; 
anterior margin broadly rounded, the carina extending nearly to lat- 
eral carinae; disk slightly flattened at middle, transversely deflexed 
along base ; surface densely, coarsely ocellate-punctate at middle, more 
densely punctate and rugose toward sides. 

Elytra moderately convex; sides parallel from bases to behind 
middle, then arcuately narrowed to the tips, which are conjointly 
broadly rounded and terminating into a short spine; surface feebly 
striate, finely rugose, rather densely, coarsely punctate basally, becom- 
ing coarsely asperate toward apices. 



NEW EUCNEMIDID BEETLES — FISHER 89 

Abdomen beneath finely, sparsely punctate; last sternite coarsely, 
densely punctate at apex, with a transversely oval, densely granulose 
area. Prosternuni, mesosternuni, and nietasternum rather densely, 
coarsely ocellate-punctate; antennal groove absent but replaced ante- 
riorly by a smooth, shining space. Metasternal episternum narrow, 
sides parallel. Posterior coxae large, strongly, triangularly expanded 
internally. 

Length 4.25 mm., width 1.25 mm. 

Type locaTit}/. — Cayamas, Cuba. 

7V;>e.— U.S.N.M. No. 57171. 

RcmwA'-f. — Described from a single male collected at the type local- 
ity, March 6, by E. A. Schwarz. 

This species is allied to Arrhipis lanieri Guerin but differs from that 
species in being smaller and of a pale reddisli-brown color, and in hav- 
ing the hairs on the elytra shorter, denser, and more uniformly dis- 
tributed, the outer segments of the antenna compact and as wide as 
long, and the prosternum flattened in front with the prosternal proc- 
ess triangular between the anterior coxae and knife-shaped behind the 
coxae. 

ARRHIPIS INSULARIS, new species 

Elongate, subcylindrical, moderately convex, strongly shining, uni- 
formly yellowish brown, antennae and legs slightly paler, sparsely 
clothed witli very short, recumbent, inconspicuous hairs. 

Head slightly convex, without depressions or carina, coarsely, 
rather densely, but not deeply, ocellate-punctate; interocular carina 
vaguely elevated, extending around inner margin of antennal fossa; 
clypeus shoit, at base twice as wide as distance to eye, broadly rounded 
in front. Antenna moniliform, slightly thickened toward apex, ex- 
tending slightly beyond base of pronotum; segment 3 nearly as long 
as following two segments united ; segments 4 to 10 vaguely wider than 
long, suboqual in length to one another; segment 11 oblong, narrowly 
rounded at apex. 

Pronf)tum slightly wider than long, widest near apex; sides nearly 
))aiallel; anterior margin broadly rounded, the carina extending 
nearly to lateral margins; disk convex, not flattened at middle, 
arcuately dcflexed along base; surface coarsely, rather densely ocel- 
late-punctate at middle, more conflucntly punctate toward si(h's. inter- 
vals vaguely granulose. 

Elytra modei'ately con\c\; sides j);ii!iIK'l fioni bases to behind 
middle, then arcuatx^ly narrowed to the tips, which are conjointly 
broadly rounded nnd terminating into 'i short spine; surface not 
striate, coarsely, rather d<'nsely jiunctate, slightly asperate toward 
apices, the intervals vaguely granulose. 

Abdomen beneath sparsely, very coarsely puncUite; last sternite 
with a transversely oval, densely granulose area. Prf)stermini. meso- 



90 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

sternum, and metasternum rather densely, coarsely punctate ; antennal 
groove absent but replaced by a smooth, shining space. Metasternal 
episternum very narrow, wider behind than in front. Posterior coxae 
not strongly expanded internally. 

Length 3 mm., width 0.9 mm. 

Type locality. — Cayamas, Cuba. 

Ty^e.— U.S.N.M. No. 67172. 

Remarks. — Described from a single male collected at the type lo- 
cality, March 15, by E. A. Schwarz. 

This species is allied to Ai-rhifis cubcmus Fisher but differs from 
that species in being yellowish brown, with very short, inconspicuous 
hairs, and in having the pronotum convex at the middle and not 
flattened, the outer segments of the antenna vaguely wider than long, 
the metasternal episternum very narrow, and wider behind than in 
front, the prosternal process triangular behind the anterior coxae, the 
pronotum arcuately deflexed along the base, and in not having the 
elytra striate. 

Genus DIRHAGUS Laporte 

DIRHAGUS ALBOFASCIATUS, new species 

Elongate, subcylindrical, moderately shining, reddish brown, except 
legs, which are yellowish brown, the pronotum and elytra ornamented 
with whitish pubescence. 

Head strongly convex, vaguely depressed near clypeus, without 
longitudinal carina, densely, finely granulose, rather densely clothed 
with short, erect, inconspicuous, whitish hairs; interocular carina 
elevated, broadly interrupted at middle; clypeus at base one-half as 
wide as distance to eye, truncate at apex; eyes large, not incised 
beneath. Antenna about two-thirds as long as body, strongly pecti- 
nate from segment 4 in male, acutely serrate from segment 3, except 
segment 11, which is narrowly elongate, in female; segment 3 much 
longer than 2. 

Pronotum distinctly wider than long, strongly convex, a short, 
longitudinal, antescutellar carina with a slight depression on each 
side ; sides parallel ; anterior margin arcuately rounded, the anterior 
supplementary carina strongly elevated, short, not extending to middle 
of pronotum ; surface densely, finely ocellate-punctate, sparsely clothed 
at middle with short, erect, inconspicuous hairs, and at sides and 
toward base with long, recumbent, whitish hairs. 

Elytra moderately convex ; sides slightly converging from bases to 
tips, which are very broadly conjointly rounded; surface vaguely 
striate along sutural margins near apices, densely, coarsely, rugosely 
punctate, sparsely clothed with short, semierect, inconspicuous hairs, 
and irregularly clothed with longer, recumbent whitish hairs along 



NEW EUCNEMIDID BEETLES — FISHER 91 

buses and skies uiul forinin<r a more or less distinct broad, transverse 
fascia behind middle and a broad spot at apices. 

Body beneath coai*sely, densely punctate, sparsely clothed with 
short, recumbent, whitish hairs; juxtasutural jjroove broad, smooth, 
sharply limited externally; lateral marj^inal carina sinuate, obsolete 
posteriorly and curving toward posterior supplenlentary carina, which 
i< stronuly elevated posteriorly, becoming: obsolete in front of middle; 
metasternal episternum narrow, wider behind than in front; posterior 
coxae slightly expanded internally. 

Length 3 mm., width 1 mm. 

Type locality. — Felton, Cuba. 

Type^ allotype., aiid paratypes. — U.S.N.M. No. 57173. 

Remnrl-a. — Described from five specimens (1 male typo). The type 
and allotype were collected at the type locality by W. M. Mann; two 
paratypes were collected at Baracoa, Cuba, during August 1901. by 
August Busck; and one paratype was collected at Higueral, Domini- 
can Republic, during February 1916, by E. G. Smyth. 

This species is allied to Dirhagus pectinatus LeConte but differs 
from that species in being smaller and in having the pubescence on the 
pr(;notum and elytra longer and ornamented with more or less dis- 
tinct white pubescent designs, the juxtasutural groove smooth, with 
the sides parallel, the posterior supplementary carina extending to 
the middle of the pronotum, the antenna in the male pectinate from 
the fourth segment, and in not having the head longitudinally carinate 
on the occiput. 

HYLOTASTELLA, new genus 

Body elongate, parallel. Head large, strongly convex; labrum in- 
visible; eyes partially covered by prot borax; clypeus strongly con- 
tracted at base. Antenna long, robust, cylindrical, segments compact ; 
second segment very small, the following segment much longer than 
wide. Pronottmi wider than long, lateral margins distinct, entire; 
anterior margin simple, not carinate. Scutellum subtriangidar, 
longer than wide, narrowly truncate at apex. Propleural triangle 
longer than wide, without antcnnal grooves. Marginal carina and 
prosternal suture of pro.sternum converging anteriorly, prostenial su- 
ture closed, distinct for entire length. Motastermun and abdomen 
without tai-sal grooves. Metastenial cpimcron completely covcicd. 
Metasternal episternum nairow. sides jiaiallel. \j\\^\ abdoininal ter- 
gite not extending over last alxlomiiial sternite. which is nai'rowlv 
rounded at ajn-x. Posterior coxa smldenly exjiaiided internally, very 
narrow externally. Logs slendei-. Tai>i withoni laniellae; segment 
1 of posterior pair as long as following segments unile(l; segment 4 
hollowed out on doi-sal surface for receiving following segment, trun 
cate at apex. 



92 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

Genotype. — HylotasteUa schwarsi, new species. 

This genus resembles a small species of Hylotastes Bonvouloir, but 
it differs from that genus in being gradually narrowed posteriorly 
and in having the antenna cylindrical and four-fifths as long as the 
body, the clypeus broadly rounded in front, the posterior coxa 
abruptly expanded internally, the propleural triangle longer than 
wide, the tips of the elytra conjointly broadly rounded, and the last 
abdominal tergite not extending over the last abdominal sternite. 

HYLOTASTELLA SCHWARZI, new species 

Narrowly elongate, moderately convex, dark brown, slightly paler 
beneath, with the humeral angles of elytra and prosternum yellowish ; 
body above subopaque, beneath slightly shining ; head and pronotmn 
densely clothed with long, recumbent, golden-yellow hairs, the hairs 
shorter and less conspicuous on elytra and underside of body. 

Head strongly convex, without depressions or longitudinal carina, 
finely, confluently ocellate-punctate ; interocular carina strongly ele- 
vated, interrupted at middle ; clypeus at base one-half as wide as dis- 
tance between it and eye, broadly, sinuately rounded at apex. An- 
tenna four-fifths as long as body ; segments 3 to 5 subequal in length 
to one another, the following segments becoming gradually longer 
to tip of antenna. 

Pronotum distinctly wider than long; sides parallel posteriorly, 
arcuately narrowed anteriorly; disk strongly convex, transversely, 
narrowly depressed along base, with a distinct round depression on 
each side in front of middle; surface finely, confluently ocellate- 
punctate similarly as on head. 

Elytra with sides parallel to apical fourth, then arcuately narrowed 
to the tips, which are conjointly broadly rounded; surface finely, dis- 
tinctly striate, the intervals finely, densely granulose. 

Body beneath very finely, densely punctate ; last abdominal sternite 
slightly gibbose and coarsely granulose toward apex. 

Length 5.75-7 mm., width 1.5-2 mm. 

Type locality. — Portobelo, Panama. 

Type and paraiypes. — U.S.N.M. No. 57174. 

Remarks. — Described from three specimens (1 type) collected at 
the type locality, the type and one paratype collected March 1, 1911, 
by E. A. Schwarz, and one paratype collected February 20, 1911, by 
August Busck. 

Genus NEMATODES Berthold 

NEMATODES EXIGUUS, new species 

Elongate, subcylindrical, moderately convex, strongly attenuate 
posteriorly, slightly shining, dark brown, nearly black, except legs 
and antennae, which are reddish brown, rather densely, uniformly 



NEW EUCNEMIDID BEETLES FISHER 93 

clothed with short, recumbent, yollowisli hairs, which arc finer on 
underside of body. 

Head rather strongly convex, vaguely depressed near clypcus, 
coarsely, densely ocellate-punctate; intcrocular carina scarcely ele- 
vated, not extending along inner margin of antennal depression; 
clypeus slightly narrower at basa than distance to eye, strongly sinuate 
at apex. Antenna robust, slightly expanded toward apex, extending 
to base of elytra; segment '2 slightly longer than 4; segment 3 longer 
than 6 and subequal in length to 4 and 5 united ; segments 7 to 9 slightly 
longer than wide : segment 11 oblong, acute at apex. 

Pronotum slightly wider than long, widest near middle; sides 
vaguely narrowed posteriorly, arcuately narrow^ed anteriorly; pos- 
terior angles acute, not carinate or divergent; disk convex, with a 
vague, round dei/i-ession on each side in front of middle, sometimes 
without depressions, rarely with a longitudinal median, smooth space; 
surface densely, finely punctate on median part, finely rugose toward 
sides. 

Elytra moderately convex; sides nearly parallel along basal half, 
strongly attenuate po^ttn-iorly to the tips, which are rather broadly, 
conjointly rounded; surface vaguely striate, the sutural stria more 
distinct, intervals flat, densely, finely punctate basally, the punctures 
becoming sparser toward apices. 

Body beneath finely, densely punctate, the punctures coarser on pro- 
sternum ; antennal groove broad, shallow posteriorly, deeper anteriorly, 
not limited internally by a carina, but sometimes with a short carina 
at outer margin near eye. 

Length 3.5-G mm., width 0.8-1.75 mm. 

Type locdJity. — Cayamas, Cuba. 

Type and paratypca. — U.8.N.M. No. 57175. 

Remcn'ks. — Described from seven specimens (1 type) collected at the 
type locality during PVbruary and March, by E. A. Schwarz. 

This species is allied to Nnnatodes alropon (Say) but dilFcrs from 
that s|)ecies in being smaller and in having the pronotum widci- Ihan 
lung, with the surface densely, finely punclate at the middle, but witli- 
out a longitudinal, median depressed line, and flic elylia vaguely 
striate. 



0. 1. soviNNiiiNr miNriNa orricc > i<49 



PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 




SMITHSONIAN INSTITUTION 

U. S. NATIONAL MUSEUM 



Vol. % Wwhington: 1945 No. 3189 

NEW LANTERNFLTES (FULGOEOIDEA) FROM 
SOUTH AMERICA 



By R. G. Fennah 



Descriptioxs are ^iven herein of seven new species of Fulgoroidea 
in tlie cixiid genus Pintalia Stal, of a new genus and species in the 
otiocerine Derbidae, and of two new species of Ateson Metcalf 
(Achilidae). All the specimens mentioned, including the types, are 
in the United States National Museum. 

Family CIXHDAE 

Genus PINTALIA Stil 

Pintalia St.\i.. Svcii.ska Vet.-Akad. IImikII., vol. 3, No. 6, p. 4, 1862. (Genotjrpe, 
P. lateralis St&l, dt'sigiiated by Muir, Pnn-I'acific Ent., vol. 1, pp. 103, 100, 
1025.) 

Tliis rapidly expanding genus is very compact. The characters by 
which species may be distinguished include the shape of the carinae 
of the vertex and of the margins of the frons at its base, that of the 
anal segment of the male, of the pygofer, aodeagus, and genital styles, 
and the color pattern on the tegmina. The natural relationship of 
the included species cannot be as.sessed with precision until there is 
evidence to indicate the order of seniority of the variables used in their 
separation. On current evidence it would .seem that the .shape of the 
male anal .segment and of the genital styles is .slightly inore stable than 
the other characters mentioned, and tiie writer tentatively grouj)S as 
follows those species for which the relevant data are availal)le: 

95 

620240 — 45 



96 PROCEEDINGS OF THE NATIONAL MUSEUM vol. w 

Anal segment short, lateroapical lobes narrow, decurved through 130°, genital 
styles short, subangulate propna group 

Anal segment fairly short, not or scarcely deflexed distally, genital styles expanded 
and spatulate distally ecuadoriensis group 

Anal segment long, distally deflexed as far as, but not exceeding, 45°, genital 
styles long, narrow in ventral view albolineata group 

Anal segment asymmetrical, often twisted into a single vertical lamina distally, 
genital styles short obtorta group 

Anal segment as above, genital styles long infuscata group 

Anal segment distally deflexed as far as 45°, genital styles short, clublike. 

oiTiata group 

Anal segment shortly deflexed through 45°, styles short, angulate, pointed in ven- 
tral view huigrensis group 

Anal segment deflexed through 90° or slightly more, genital styles distally ex- 
panded, truncate or concave on apical margin Mcaudata group 

Anal segment as above, very asymmetrical, genital styles rounded at apical 
margin vomerifera group 

The propria group includes only this species; the second group 
includes ecuadoriensis Muir and quadrimaculata (described below) ; 
albolineata Muir, by contrast, typifies a large group including hnm- 
nivenosa Muir, latinotata Muir, longispinis Muir, quadrispinosa Muir, 
fuscomaculata Muir, hlairmontensis Muir, tumatumariensis Muir, 
fuscipennis Muir, alhomarginata Muir, ohscurata Muir, and two 
species, raamiorata and ohliquivitta^ described below; the ohtorta 
group, including obtorta Muir, pulchella Muir, and fuscomarginata 
Muir, would appear to be quite closely related to the infuscata group 
{infuscata Muir, angustinofata Muir, and falcata (described below) ) 
if judged by the shape of the anal segment; omata Muir is grouped 
with altamasonica Muir and discrepans Muir, huigrensis Muir is 
somewhat doubtfully associated with furcata Muir, while bicaudata 
Muir, maculipennis Muir, and curvivitta and daedala (both described 
below) form a fairly closely knit group ; vomerifera (described below) 
is placed apart, though it shares characters with the preceding group 
and with the obtorta group, on the ground that the shape of the genital 
styles and the general structure of the aedeagus differ considerably 
from those of all the species so far considered. 

PINTALIA QUADRIMACULATA, new species 
PlATE 2, FiGUBES 1, 7-10 

Male. — Length, 4.0 mm. ; tegmen, 5.9 mm. 

. Vertex with anterior margin and transverse carina straight, lateral 
margins of f rons not thickened in basal portion. 

Fuscous ; genae, lateral fields of pronotum, sternum, and legs testa- 
ceous. Tegmina testaceous, posterior edge of costal cell, base of cell 
Sc, basal four-fifths of cell R, cell M, and the larger part of cells Cu^a 
and Cuib deeply inf uscate, almost piceous, a white round spot at middle 



NEW LANTERNFLIES FROM SOUTH AMERICA — FENNAH 97 

of common stalk of M, a similar spot at fork of Cui, a third on Cuia 
and a fourth on Cuib, a hyaline area parallel to nodal line and just 
basad of it, a fuscous band overlying nodal line and a similar band of 
equal width and more clearh' defined obliquely traversing apical cells 
acro:ss middle from apex of K to M3, apical margin narrowly fuscous 
to M3 then broadly fuscous posteriorly but infuscate area interrupted 
by a large testaceous spot distad of apex of clavus. 

Anal segment of male short, not deflexed; lateroapical angles only 
slightly prominent, very slightly produced ventrally. Pygofer broad 
■with each lateral angle produced in a short narrow lobe, distally 
rounded. Aedeagus tubular, straight, with three spines arising at 
approximately same level at base of flagellum, the middle spine por- 
rect, the other two strongly curved. Genital styles in profile angulato 
at middle with a pointed eminence on dorsal border, spatulate distally 
with greatest width two-thirds from base. 

Ti/pe.—V.S.^M. No. 57092. 

Described from one male specimen taken at Tumupasa, Bolivia, by 
W. M. Mann (December 1921). The dark basal field of the tegmina 
and its ornamentation of four cretaceous spots readily distinguish 
this species. 

PINTALIA MARMORATA, new species 

Plate 2, Fiqubes 5, 20 ; Plate 3, FiGtTBES 30-32 

Male. — Length, 4.1 mm. ; tegmen, 6.0 mm. 

Vertex with anterior margin and transverse carina straight, lateral 
margins of frons very slightly thickened basally. 

Testaceous; frons, genae, lateral fields of pronotum, and mesonotum 
fuscous. Tegmina ivory-hyaline, marbled dull brown in small irregu- 
lar spots, with an irregular fascia across apical cells subparallel with 
apical margin; veins dull brown. Wings hyaline, veins fuscous. 

Anal segment of male moderately long, deflexed through 45° distad 
of anal foramen but not much produced. Pygofor broad, lateral 
angles large, with dorsal margin sliglitly concave, ventral margin con- 
vex. Aedeagus long, tubular, slightly curved dorsad, with three un- 
equal spines at base of flagellum, one being much longer than the other 
two, flagellum distally expanded, its dorsal margin contiiuied in a 
slender curved spine, apex of flagellum somewhat pointed. Genital 
styles strongly angulaf o, with a broad-basod, pointed eminence dorsally 
near angle, gradually ex[)anded distally, apical margin obli(iuely 
truncate. 

Ti/pr.—V.S.yiM. No. 57093. 

Described from a single male specimen collected at light on deck 
in harbor, Puerta Bolfvar, Ecuador, by M. Kisliuk and C. E. Coolcy 
(April 9, 19:'.2). This sj^ecies is distinguished by the pattern on tho 
tegmina and by the shape of the anal segment, pygofer, and genitalia. 



98 PROCEEDINGS OF THE NATIONAL MUSEUM vol. B6 

PINT ALIA OBLIQUIVITTA, new species 
Plate 2, Figuees 3, 16-19 

Male.—Length., 4.0 mm. ; tegmen, 6.0 mm. 

Vertex with anterior margin and transverse carina straight, lateral 
margins of frons prominently raised but not thickened basally. 

Fuscous; sternum testaceous, legs pale fuscous. Tegmina ivory- 
hyaline, a broad oblique band from base of stigma to apex of clavus, 
three transverse narrow bars in costal cell, the basal bar extended in a 
narrow broken fascia to claval suture, a suffusion along nodal line 
and two S-shaped areas in apical cells of E, and M, a pale suffusion 
over distal part of apical cells, cell Cuib except for a round spot, and 
two broad bands between claval suture and first claval vein yellowish 
brown. Wings hyaline, veins testaceous. 

Anal segment of male moderately long, distally deflexed through 
60°, deflexed part not nearly so long as horizontal part. Pygofer 
broad, lateral angle about 60°. Aedeagus tubular, a spine one-third 
from base on left side directed ventrally and anteriorly, a second spine 
on left side at base of flagellum directed dorsally and caudad, a stout 
curved spine at middle of dorsal surface directed dorsally and caudad, 
a longer slender spine on right side at base of flagellum directed ven- 
trally and anteriorly, flagellum with two minute adpressed spines 
on right side near middle, and a longer somewhat oblique spine at 
apex. Genital styles angulate near base, with a short broad spine 
dorsally near angle, distally narrow, slightly expanding caudad, 
rounded and setigerous at apex. 

Type.—V.S.l^M. No. 57094. 

Described from one male taken at Santo Domingo, southeastern 
Peru, labeled "collection Kosenberg." This species is distinguished 
by the genitalia and by the tegminal pattern. 

PINTALIA FALCATA, new species 

Plate 2, Figures 4, 22 ; Plate 3, Figures 27-29 

Male. — ^Length, 4.3 mm. ; tegmen, 6.1 mm. 

Vertex with anterior margin and transverse carina straight, lateral 
margins of frons not thickened basally. 

Fuscous ; tegmina ivory-hyaline, extreme base of tegmina, a broken 
fascia from costa to junction of claval veins, a second fascia from 
costa just basad of stigma to commissural margin distad of apex of 
clavus, nodal line, apex of each apical vein, and a band from third 
apical cell of M to middle of cell Cuia yellowish brown. Wings 
slightly infuscate, veins dark. 

Anal segment of male angulately deflexed beyond anal foramen, 
twisted into a horizontal lobe distally. Pygofer with lateral angles 
broadly rounded. Aedeagus tubular, a long straight spine on left 



NEW LANTERNFLIES FROM SOUTH AMERICA — FENNAH 99 

side at base of fl:i^olIum, a lon<^ curved spine on right side at same 
level, both directed anteriorly; llagelluni with a curved tubular mem- 
branous lobo arising at middle and curving to left, a pair of broad 
unequal spines distally. Genital styles weakly angulate at middle, 
distal portion somewhat sinuately expanded, a row of minute denticles 
on inner border distally. 

Type and parafr/pe.—V.S.^M. No. 57095. 

Described from one male and one female collected in French Guiana 
by W. M. Schaus. 

PINTALIA CURVrVITTA. new species 

Plate 2, Figures 2, 11-15 

Malr. — Length, 3.9 nmi,; tegmen, 5.9 mm. 

Vertex with anterior margin and transverse carina straight, lateral 
margins of frons not thickened basally. 

Fuscous; sternum and legs testaceous. Teginina ivory-hyaline, 
sparsely marked with yellowish brown on membrane, a broad approxi- 
mately semicircular band of the same color widely enclosing stigma 
and reaching posteriorly to Cuia; a band across apical cells of M, apex 
of Cuib, a spot distad of apex of clavus, and a suffusion at middle 
of claval suture yellowish brown. "Wings hyaline, veins testaceous. 

Anal segment of male deflexed through 100°, symmetrical, deflexed 
part almost as long as basal part. P3'gofer with lateral angles pre- 
duced into a short narrow lobe, markedly sinuate at apex. Aedcagus 
tubular, a short stout spine on dorsal surface one quarter from base, 
directed dorsally and unti-riorly, a second spine of similar shape, 
though slenderer, dorsally at middle, and a third spine, curved dorsad 
and caudad, at base of flagollum, a short slender spine at middle on 
left side directed ventrad and forward, a longer sinuate spine sub- 
parallel to it at base of flagellum, flagellum devoid of ornamentation. 
Genital styles angulatcly bent at middle, expanded distally, apical 
margin very shallowdy excavate. 

rv/?e.— U.S.N.M. No. 57096. 

D('-<ri!)cd from a single male taken at Rurrenabaque, Bolivia, by 
W. M. Mann (November 1921). This species is distinguished by tho 
shaiK' of the anal segment, of the lateral angles of the pygofer and of 
the genitalia, and by the pattern on tho tegmina. 

PINTALIA DAEDALA, new iipcrica 
Pi>ATE 2, FioxjRKS n, 21 ; Pi.Ai>: .3, Fi<;ure8 33-35 

Male. — length, 5.0 mm.; tegmen, 0.0 mm. 

Vertex with anterior margin straight or slightly curved, transverse 
carina slightly arcuate on each side of middle line, lateral margins of 
frons thickened basally. 



100 PROCEEDINGS OF THE NATIONAL MUSEUM vol. M 

Anterior portion of vertex, basal three-quarters of f rons, distal por- 
tion of clypeus, sides of clypeus and genae before eyes, pronotum 
medially, mesonotum, mesopleurites, and fore and middle legs piceous- 
brown ; posterior portion of vertex, f rons in apical quarter and clypeus 
at base, genae above eyes and around antennae, lateral fields of prono- 
tum and basal half of procoxae ivory to pallid, hind legs and sternum 
pallid testaceous, abdomen dull fuscous. Tegmina hyaline, basal two- 
thirds of clavus yellowish, postcubital (first claval) vein dark in middle, 
a yellowish-brown fascia from two bars at middle of costal cell to com- 
missural margin at level of union of claval veins, a very broad fuscous 
band transversely across tegmina, enclosing stigma anteriorly and of 
subequal width throughout, a narrow wedge-shaped band from apical 
veins of R to penultimate apical cell of M, apical margin expandingly 
fuscous from Rs, two opalescent-hyaline spots between stigma and apex 
of clavus between Rs and M and Mo and M3+4, respectively, the veins 
at Mf , M-Cu, and Cu distad of apex of clavus yellow. Wings smoky, 
veins testaceous. 

Anal segment of male deflexed distally through 80°, deflexed portion 
fully as long as basal, in profile slightly dilated distally, a semi-cir- 
cular excavation on apical margin at middle. Pygofer with lateral 
angles shortly produced in a small rounded lobe. Aedeagus tubular, 
a short, broad-based horizontal spine at middle on right side, and a 
slenderer spine directed caudad at same level on left side, two long' 
spines at base of flagellum on left side, one curved dorsad, the other 
sinuately ventrocaudad, flagellum tubular, somewhat tumid at base. 

Type.—V.S.'NM. No. 57097. 

Described from one male taken near Bailos, Ecuador, by S. W. 
Frost (February 20, 1937) . This species is distinguished by the shape 
of the anal segment of the male and of the genitalia and by the pattern 
on the tegmina. 

PINTALL4 VOMERIFERA, new species 

Plate 3, Figures 36-38 

Male. — ^Length, 4.7 mm. ; tegmen, 5.5 mm. 

Vertex with apical margin and transverse carina straight, lateral 
margins of f rons not thickened basally. 

Testaceous- fuscous ; tegmina ivory-hyaline, three brown spots in 
costal cell, the basal spot extending in a broken fascia faintly across to 
junction of claval veins, a brown spot over R-M and M-Cu, membrane 
slightly and unevenly suffused fuscous, vein Cuib pallid, a pale spot in 
middle of cell Cuia. Wings very slightly smoky, veins testaceous. 

Anal segment of male deflexed distally through 95°, right side 
dilated, apical margin very oblique, making apex acutely pointed. 
Pygofer with lateral angles not produced, or, if so, very obtusely. 



NEW LANTERNFLIES FROM SOUTH AMERICA — FENNAH 101 

Aedea^^is tubular, a stout spine in middle line ventrally curved pos- 
teriorly, a deep median keellike lobe ventrally in distal half, a long 
bladelike process arising at base of flagellum on right directed ventrally 
and forwanl, two unequal spines at base of flagellum on distal side 
directed dorsally, flagellum with two scroll-like folds. Genital styles 
weakly angulate at middle, much dilated distally, outer margins 
strongly convex, inner margin straight or nearly so. 

Typc—V.S.l^M. No. 57098. 

Described from one male taken at Banos, Ecuador, by S. W. Frost 
(February 20, 1937). This species is distinguished by the shape of the 
anal segment and of the genitalia and by the pattern on the tegmina. 

Family DERBIDAE 

Tribe OTIOCERINI 
IQUITOSA, new genus 

Vertex longer than wide across base (3 :1) , disk deeply sunken, lateral 
margins thickened and pustulate distally, approximated at apex, pos- 
terior margin concave; frons linear, clypeus medially carinate; head 
compressed, in profile produced obliquely upward and anteriorly in an 
acute angle, almost pointed at tip ; antennae subequal to length of head, 
first joint about as broad as long, second joint flattened, with sides 
straight, expanding distally, apical margin asj'mmetrically excavate, a 
single short vermiform appendage attached at base. Pronotum very 
short, anterior margin angularly convex, posterior margin still more 
acutely concave, lateral fields broad, quadrate, median carina present, 
and a carina on each side between eye and tegula ; me«onotum convex, 
depressed in posterior third, median carina feeble, lateral carinae obso- 
lete except for two flangelike vertical eminences in middle; tegulae rel- 
atively large. Tegmina with sides expanding distally, apical margin 
oblique, costal margin sinuat<^, with a prominent recurved rounded 
eminence near base, vein M leaving Sc + R one-seventh from base, 
Sc + R forking two-sevenths from base, M with five branches reach- 
ing apical line of transverse veins, clavus narrowly open, but common 
claval vein not passing beyond its apex. Wings four-fifths as long as 
tegmina. 

Genotype, Iquttoaa shannoiu^ new species. 

IQL'ITGSA SHANNONI. new Hprrin 

PI.ATC 2. FI0URE8 23-26; Platk :{. Fkjukes 31M1 

MaJe. — Length, 3.2 nnn. ; togmen, 5.0 mm. 

Head, pronotum except margins, mesonotum except carina, a patch 
on each lateral field, and posterior margin brown; median carina of 
mesonotum and margins of pron(*tum and mesonotum, sternite^s, and 



102 PROCEEDINGS OF THE NATIONAL MUSEUM vol. w 

legs testaceous to stramineous, except for a narrow fuscous band at 
apex of profemora and a similar band in middle of protibiae. Teg- 
mina mostly piceous, a dull testaceous spot on each side of piceous 
costal eminence, apical cells from node to Ms and intervening veins dull 
yellow with two fuscous interruptions, middle portion of subapical 
veins paler than corium, veins of corium chiefly red, basally infuscate 
and interrupted with dull yellow spots. Wings smoky, veins con- 
colorous, margin red. 

Anal segment narrow, lateral apical angles slightly deflexed. 
Aedeagus tubular in basal half, with a minute bicuspidate process 
near base of flagellum, two stout spines at base of flagellum, one on 
right side, the other distally, both directed dorsally, a transparent 
spine on flagellum at middle, apex of flagellum lobate with a large 
spine below it on left ; two small spines on right. Genital styles nar- 
row, membranous dorsally distad of middle, with a scroll-like process 
in middle of dorsal margin. 

Type.—U.S.^M. No. 57099. 

Described from one male collected at Iquitos, Peru, by R. C. Shan- 
non (March 1931). The basal reflection of the costal margin recalls 
the similar modification found in Sayiana Ball, but Iquitosa differs 
from Ball's type, with which it has been compared, in the more pointed 
head, the shape of the antennae, and the shape and venation of the 
tegmina. 

Family ACHILIDAE 

Subfamily APATESONINAE 

Genus ATESON Metcalf 

Ateson Metcalf, Bull. Mus. Comp. Zool., vol. 82, No. 5, pp. 367, 369, 1938. (Geno- 
type, A. marmoratum Metcalf, loc. cit., p. 369.) 

ATESON SEMILUTEUM, newr species 

Plate 3, Figures 42-47 

Male. — ^Length, 5.5 mm. ; tegmen, 8.0 mm. 

Frons with lateral margins ampliate and raised between level of 
antennae and suture, median carina terminating distally against a 
transverse ridge, median carina of clypeus distinct except at base. 

Testaceous ; abdominal sclerites infuscate. Tegmina yellowish, fus- 
cous at base and chiefly so distad of nodal line, with a pale area in the 
subapical cells of M and Cuia, transA^erse veins mostly pale. Wings 
hyaline at base, smoky near apical margin, veins fuscous. 

Anal segment of male short. Pygofer with lateral angles scarcely 
produced, pointed, slightly falcate, medioventral process twice as 
broad as long, distally shallowly trilobed, the middle lobe largest. 



NEW LAXTERXFLIES FROM SOUTH AMERICA — FEXXAH 103 

Aedeagus tubular, sli^litly taporiun; distally, pcriandiium rocurved 
dorsally at apex into a sinuate spine, penis when everted with two 
curved spines directed ventrally and two straight spines porrect 
caudad. Genital styles distally expanded, in profile with apical mar- 
gin sinuately oblique, dorsal margin sinuately horizontal, folding in- 
ward with a short pointed lobe. 

Tijpe.—JJ.SSM No. 57100. 

Described from one male taken at Para, Brazil, by P. R. Uhler 
(coll. No. 132). This species is distinguished by the carination of 
the frons and by the pattern on the tegmina, as well as by the shape of 
the genitalia. 

ATESON LUTEOSPERSUM, new species 

Plate 3, Fiouurs 48, 49 

Fe77iaJe. — Length, 7.0 mm. ; tegmen, 8.5 mm. 

Lateral margins of frons not much raised above level of disk, disk 
of vertex scarcely depressed, margins scarcely raised. 

Vertex, frons, clypeus, pronotum, and mesonotum fuscous, heavily 
speckled with yellow spots, genae testaceous, sternum and legs fuscous. 
Tegmina fuscous, heavily marbled and spotted with yellow on mem- 
brane, a small patch basad of stigma and over junction of claval veins 
yellow, corium wholly fuscous, veins spotted with yellow. Wings 
smoky, veins fuscous. 

Ovipositor with first valvulae bearing two teeth dorsally near apex 
and a curved apical spine; apex of second valvulae rounded in profile. 
Genital chamber (bursa copulatrix) with three pairs of sclerotized 
plat<'.s, each of the posterior pair, in dorsal view, elongate-quadrate, 
each of the middle pair larger, curved and triangular, each of the 
third (dorsal) pair small, narrow, sinuately tapering as shown in 
figure. 

7y/>c.— U.S.N.M. No. 57101. 

Descril)ed from one female taken at Cabima, Panama, by A. Busck 
(May 28, 1911). Thi.s species is distinguished by the weak carination 
of the frons and vertex, by the shape of the second valvulae of the 
ovipositor and of the sclerites of the genital chamber, and by tho 
distribution of the yellow spots on the tegmina. 

ATESON MARMORATUM Metralf 
Pr.ATi: 3, FifiUKES r»0-54 

AtCHon mannoralum MfrrcAu. Bull. Miih. Coinp. Z<)oI., vol. 82, No. 5, p. 300, 1938. 

For comparison with the preceding species figures are given of the 
male genitalia of a paratype specimen and of tho distinctive portions 
of the female genitalia. 



EXPLANATION OF PLATES 

Plate 2 

1,7-10, Pintalta quadrimaculata, new species: 1, Tegmen; 7, vertex; 8, anal segment, 
pygofer, and right genital style; 9, aedeagus, right side; 10, aedeagus, left side. 
2, 11-15, Pintalia curvivitta, new species: 2, Tegmen; 11, vertex; 12, anal segment and 
pygofer; 13, right genital style; 14, aedeagus, left side; IS, aedeagus, right side. 
3, 16-19, Pintalia obliquivitta, new species: 3, Tegmen; 16, vertex; 17, anal segment, pygofer, 
and right genital style; 18, aedeagus, right side; 19, aedeagus, left side. 
4,22, Pintalia falcata, new species: 4, Tegmen; 22, anal segment, pygofer, and right 
genital style. 

5, 20, Pintalia marmorata, new species: 5, Tegmen; 20, vertex. 

6, 21, Pintalia daedala, new species: 6, Tegmen; 21, vertex. 

23-26, Iguitosa shannoni, new genus and species: 23, Anal segment, side view; 24, 
aedeagus, left side; 25, aedeagus, right side; 26, right genital style, side view. 



Plate 3 

27-29, Pintalia falcata, new species: 27, Anal segment, left side; 28, aedeagus, left side; 
29, aedeagus, right side. 

30-32, Pintalia marmorata, new species: 30, Anal segment, pygofer, and right genital 
style; 31, aedeagus, right side; 32, aedeagus, left side. 

33-35, Pintalia daedala, new species: 33, Anal segment, pygofer, and right genital style; 
34, aedeagus^ right side; 35, aedeagus, left side. 

36-38, Pintalia vomerifera, new species: 36, Anal segment, posterior view; 37, anal seg- 
ment, pygofer, and left genital style; 38, aedeagus, right side. 

39-41, Iquitosa shannoni, new genus and species: 39, Head and pronotum (right antenna 
incomplete); 40, head in profile; 41, tegmen. 

42-47, Ateson semiluteum, new species: 42, Anal segment and pygofer; 43, medioventral 
process of pygofer; 44, aedeagus (everted), right side; 45, right genital style; 
46, head, frontal view; 47, tegmen. 

48,49, Ateson luteospersum, new species: 48, Right second valvula of ovipositor; 49, 
sclerotization of genital chamber. 

50-54, Ateson marmoratum Metcalf : 50, Right second valvula of ovipositor; 51, sclerotiza- 
tion of genital chamber; 52, right genital style, ventrolateral view; 53, medio- 
ventral process of pygofer; 54, aedeagus (retracted), left side. 

104 



U. S. NATIONAL MUSEUM 



PROCEEDINGS. VOL. 96 PLATE 2 




NEW South American Fulgoroidea, 
For explanation 8cc page 104, 



U. S. NATIONAL MUSEUM 



PROCEEDINGS. VOL. 96 PLATE 3 
33 




South American Fulgoroidea. 
For explanation see page 104. 



U. S. GOVERNMENT PRINTINS OFFICE: I94B 



PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 



h the 



SMITHSONIAN INSTITUTION 
IS NATIONAL MUSEUM 




Vol. 96 Washington: 1945 No. 3190 



TTIF. OKXT'S FT'XDELLA ZELLKK: A COXTRTBrTIOX 
low AIM) A IJFAISIOX OF THE AMEKTCAX PVKALI- 
DoID Moril-^ OF rilF F.V.MILY PHYCl'lTD.VF 



B\- (".\i;r. IIi;im;ii ir 



This paper i.s oftVred now because two of the species treated liave 
iMcn discovered in the United States, and one of these {pellurcnf<) is 
rated as a l)ean pest of some importance in the Tropics and may easily 
become one in our Gulf States. Also, there has been considerable mis- 
identificaHon of at least three of the species due to the wide variability 
of the markings and the mi>ai)i)lication of the name peUucens in eco- 
nomic and taxonomic literal iin-. Tlic >])('(ies are markedly dilfeicnt in 
theii" genitalia, but herelofoic tlic'-e organs have not l)een figured, nor 
have the structuial iiecnliaril ies of the several species been indicated. 

The genus is conliii'il \n ilic Niw A\'(irl(|. h'ivc species nrc recog- 
nized of which one is iier<' described for tiie jii-.^t time. Two older 
names an; reft ri'ed to synonymy. T'lie >l udy is l);i<e(l upon material in 
the United Slates National Museum, a considcr;il>le collect imi id' speci- 
mens from Puerto Kieo in the Cornell Uidxcrsity collection, and a few 
s|iecimens from the .Janse and ]*riti~h .Museum collections. 

(Jcmis II NDKI.I.A /»ll.r 

rmiili llii Zki.i.kk, I.sIs voii (»kfii, \(»1. 11, p. stu;, l.SJs,— U.vi.onoi', .Mciiioircs sur le.s 
I/-pi(l(iI(terf'S, vol. 7, p. UlU. lW.i. — .Iansk., Jeiirii. Kiit. Soc. Soiilli Africa, vol. 
•I, p. 1W. 1041. ((Jennfyi)o: Fundrlla jkUuciiik Zellcr.) 

lUillnvin DvAU. I'rr»c. U. S. Niif. Mils., vol. 41, p. '.VS.\, VM'.\: Iii.sociilor Tiisciii.ic 
Mfiistnuis, vol, 7, p. in, UM'.*. («;i'iintyp<' : Itnllnrin cisHin iiiiis Dyar.) 

Anleima of male slightly piibe.-cent, base somewhat enlarged, shaft 
laterally (lattened ami very slightly excavate at ba.se (pi. 4, fig. li) 

•'.2024.". 4.'i in.'. 



106 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

(except in k/nohUis and aliemora) and -with a very small blackish scale 
tuft in the excavation (except in ignohilis) ; of female slender, simple. 
Front of male head deeply grooved to hold labial palpi ; of female 
rounded. Lal;ial palpus upcurved, reaching to vertex, clothed with 
broad appressed scales; in male closely appressed to face, with second 
segment over three times as long as first and with third segment very 
short (about one-sixth the len.gth of second) ; in female with second 
segment shorter and third about one-third the length of second. Max- 
illary palpus minute. Forewirig smooth ; 11 veins, 10 from cell, parallel 
for some distance but not approximate to stalk of 8-9, 9 short, 6 from 
below upper angle of cell, straight, 4-5 connate or ajjproximate at base, 
3 approximately equidistant from 4 and 2, 2 from before lower angle 
of cell. Hind wing with 8 veins, 7 and 8 closely approxinuite beyond 
cell for less than half their lengths, 4 and 5 long stalked, 3 from stalk 
of 4-5 or closely approximate for some distance, 2 from near lower 
angle of cell, cell short, discocellular slanting and slightly curved; in 
male anal area, involving veins la and lb, thickened and folded under 
to form a pocket enclosing enlarged scales and hair tufts. Eighth 
abdominal segment of male bearing a thin, short pair of ventiolaterai 
hair tufts. 

Male genitalia with uncus long, curved, strongly sclerotized, con- 
stricted at middle and broadly divided at apex (hammer-clawed) ; 
gnathos terminating in a short, stout hook or a short, broad plate 
{ahemora) ; harpe rather short, with clasper ; vinculum narrow, short; 
aedeagus stout w4th long, stout, projecting, curved spine or spines 
at apex (except in argent ina) ; cornutus a single, strong spine. 

Female genitalia without signum {jyellucens) or with signum well 
developed and consisting of a large oval or pear-shaped cluster of 
thornlike spines {^argentina^ agapella), or curved sclerotized bands 
armed with stout, thornlike spines {ahemora^ ignohilis) ; bursa large; 
ductus bursae short, broad (narrowest in agdfeUa) ; area surround- 
ing genital opening strongly sclerotized, the dorsal sclerotization in 
the form of a band connected with the supporting rods of eighth 
segment collar, and armed with two or four spinelike projections 
(except in ignohilis and some examples of argentlna) ; ductus semi- 
nalis from caudal area of bursa. 

This genus is easily distinguished by its striking male characters: 
the strongly sclerotized, long-stemmed, bifurcate (hammer-clawed) 
uncus; the large pocket on anal area of hind wing; the long, embedded 
labial palpus with very short third segment; and minute maxillary 
palpus. A similar bifurcate uncus is not found in any other Ameri- 
can genus except Defundella Dyar. In the type species of the latter 
{corynophora Dyar) the uncus is somewhat produced and exhibits a 
slight bifurcation at apex; but other species, which must also be 



THE (.KM - IINDKI.I.A HKIXHK II 107 

1 t'ft'iTcil to I >< fiiiuli'llii, l.itk I Ills cluu'ncii'r. I >( fumhlJa soparatos 
rt'Mflily nil otlu-r male st i"iutiirc> : Its irn-allv in I need irnatlios, j^tron^rly 
linokcil. partially free sacciiliiv of liarpc it> rniiiidcd linns, and the 
1. arrow. vti"on!_dy >-cIt'ini izt'd. deeply iii\ aeinatfd pmkfi of i he .-icriiitc 
id" tlu' I'iirhtli alulouiiiial >('i:iih'1ii . 

In Fuiiililht tilt' wiii<2' patirrii \;iric- ^-o imiili within aii\- !.M\('n 
-pt'cies tliat it atl'ofds no irlialilc chniacit'r I'or >pcfitic iih'iii ilicai ion. 
and the sm'iM'al -pccics can he -cpai-alrd with ct'ilaintx onl\- h\ thi ii- 
•ronilalia. 

KEY TU Till-: Sl'KCIKS UF l-UNDKLLA 

MALES 

]. ('l.isiM-r ;i str;ii};lii >|iiiir; m<m|.',i;:us simple argentina Dyar 

• 'laspor a curveil (li;rinis: ,h'iji":il:us aiiiiPtI with curved, strongly sclerotized 

spiii«> nr spines at or near ai»ex 2 

_' W itii larjje, stronfily sclerotized snbanal plate; a cluster of several spines 

from apex of aedeagus; <-laspei- short pellucens Zeller 

Witlmut srlerotized sut»aiial plate; no more than two spines from aedeagus 

at or near ajiex; clasper long 3 

'■\. (inatlios terminating in a l)road plate; aedeagus with a pair of spines from 

apex ahemora Dyar 

(Inathos terminating in a short, stout hook; aedeagus with a single spine 
from hel(»w ajK'X ignobilis, new siK'cies 

FKMALKS 

1. r.nrsa copnlatrix wiilii>iit slgnum : (liniiis Imrsae sclerotizi'd throughout 
leiigtli, scjerntizaliiin involving part of bursa adjacent to ductus Iiursae 

aiirl (hielns si'ininalis pellucens Zeller 

r.nrsa eopnlairix with strong signa ; ductus bursae at most only partially 

sr-lernt i/.Ml 2 

-' Simia iti fmiii nf lar^'e <>\al nr iiear-s|i,i|pei| cjuslers of spines argentina Jtyar 

agapella Schaus 

SiL'iia ill fiiriii nf bands beariiiir sloiil si)iiies 3 

'■'■. Signa eniisisiiiig of If rallier slinri bands, eaeli arnietl witli a rnw of bmg 
spines: duel lis Itiirsae willi a slrmigly seleroti/.ed median ndlar; a widely 
sjiaee*! pair of long, curved spines from sclerotized area immediately lu'liind 

geniial opening ahemora Dyar 

Sigiiji <*<>nsisling (if '1 loni:, partially fused bands eaeli ariiieil with a inw nf 
slmrl, stniit, llHtriilike sjiines ; diielns bursae with median area uiisclero- 
lize«l; no spiiM-s adjacent to genital ojM'iiing ignobilis, new si)ecles 

Kl Mil I I \ I'l II I I I NS /. ll.i 
IM.MI I. Fk.I Kl > 1 I. t; t;.(. 7 7' ; I'l Ml Ci. Kk.IKI s 1". 1."../ 

I iiii'lillii iiflhniiiM Zn I Kit. Isis vun nken, Mil. II, p. sCiC, |s)S: llmae See. lint. 
Kossieae, Vi>l. pi. p. !.•:{(;, ISSl. |{A(iOMiT. .Memnires slir les l/''lii(l"|ileres. 

vol. 7. p. 2ln, ]su:\. 
Hitll'iii)! rixtipriiuiM Dv mj. I'hk-. I'. .»<. Nat. Mils., vel. H, p. .•{::;',. P,ii:{. 



108 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

Fundella cistipennis (Dyar) Dyar, lusecutor Inscitiae Meustruus, vol. 7, p. 40, 
1919.~WoLCOTT, Jonrn. Dept. Agr. Puerto Rico, vol. 17, pp. 241-255, 1933; 
Journ. Agr. Univ. Puerto Rico, vol. 18, p. 432, 1934; vol. 20, p. 477, 1936.— 
Scott, Journ. Agr. Univ. Puerto Rico, vol. 24, pp. 35-47, 1940. (New 
synonymy. ) 

Male. — Antennal shaft with very small black basal tuft (pi. 4, 
fig. 3). Palpi, thorax, and forewing grayish fuscous more or less 
dusted with whitish and with interspersed reddish-brown scales (in 
many specimens the ground color is reddish brown), giving the moth 
a distinctly gray or gray-brown appearance to the naked eye. Fore- 
wing with a conspicuous, round, darker brown or fuscous sj)ot in 
the center of the area usually occupied by the antemedian line, this 
dark spot more or less obscured in some specimens but in typical 
examples outlined by whitish areas inwardly and outwardly and not 
reaching to inner margin or costa of the wing; discal mark at end of 
cell obscure, often absent; subterminal line (when distinguishable) 
faint, white, indented at vein 6 and at submedian fold ; a row of dark 
spots along termen (present only in specimens having an appreciable 
dusting of white scales). Hind wing white, translucent, a faint 
fuscous border along costa and (in some specimens) a fuscous line on 
termen for a short distance from apex; cilia white ; anal pocket yellow- 
ish white. Midtibia with a fringe of pale hairlike scales along dorsum. 
Hind tibia with a rather long and slender tuft of pale (whitish ochre- 
ous) , hairlike scales from the knee joint (pi. 4, fig. 4) . 

Alar expanse 19-23 mm. 

Genitalia (pi. 4, figs. 7-7c) with a large, strongly sclerotized sub- 
anal plate, constricted before and beyond its middle. Harpe with 
apex notched below costa; clasper short, curved, situated near middle 
of harpe and armed with several setae at its knobbed apex. Aedeagus 
Avith a cluster of several long, curved spines from apex ; cornutus long, 
straight, stout. 

Female. — Essentially like the male in color and markings except 
that the dark spot near the base of the forewing is more diffused, 
sometimes reaching to the costa. Hind wing usually with a dark shade 
along termen. 

Alar expanse 19-24 mm. : 

Genitalia (pi. G, figs. 15-15a) with bursa copulatrix finely scobinate 
but without signum ; ductus bursae flattened, broad, twisted and con- 
stricted near genital opening, sclerotized throughout, the sclerotiza- 
tion involving bursa adjacent to ductus bursae and ductus seminalis; 
sclerotized band behind genital opening armed with four long, stout, 
projecting spines; collar of eighth segment invaginated at dorsal 
margin to form a sclerotized pocket (pi. 6, fig. 15«). 

Types. — In British Museum {peJluce?is) ; United States National 
Museum ( cistipennis ) . 



THE GENUS FUXDELLA — HKIXIUCH 109 

Tiipe localitks. — St. Thomas-. l?iiii>li AVest Indies {pclluecn-s) ; 
B:i rl )atlos {cist i pen n As ) . 

Food plants. — Vigna un</iii<-uhita ((•(•wpi-as. l)l;i(k-fyt'(l pt'iin, and 
<:ardei\ i^cas), Cunavalia cnsiformis (sword beans), Canavalla mari- 
time (black bean), Cnjan cajan (pigeon poa), Pha'^cohis hnmtiis (cul- 
tivated ami wild lima beans), PJiascohis sp. (Brazilian specimens), 
Cassia occidcntalis (one reared siiecimcn I'mni McCubbins Mills, 
Puerto I\ico. before me; most records from this last plant are doubtful 
aiul proiiably the result of a misidentification of FundclJa argcntina as 
( istipcnnis). 

I distribution. — I'nited States: Florida. Hobe Soiuid. ^Sliami. Jupi- 
ter. Coconut Grove, Marco Island, AValton, Jensen (U.S.D.A. rearings 
Ik .in lima beans, February 1944), Vero Beach (J. R. Malloch, Decem- 
ber r.>4r). B.MMVMHis. IlArrr : Damien (Deceml)cr, Fel)ruary). Bort- 
au-Piince. Montsi:i:i;at (January). Cuba: Santiago, Matanzas. 
\'ii;(.iN Islands: St. Croix (March). Puerto Rico: San Juan, Rio 
Picclias (^larch-May), Isabella, Catano (July), Vieques Island 
(April), Bkazil: Bahia (May), Ceara. 

Xinetj'-six specimens examined. 

Zeller had two species before him when he described pcJhiccns., and 
the one he figured (fig. 411) in Horae Soc. Ent. Rossicae is his ''var. 
!»,*• which is Dvar's argcntina. Through the courtesy of Messrs. Riley 
and Tams. of the British Museum, I have been able to examine the 
genitalia of the male paraty]»o de<i^nated "var. b" by Zeller and a 
typical male pclluccns of the Zellei' material from ISIai'aiinita. The 
latter proved to be what Dyar described as cistipcnnis and the species 
that has ajipeai-ed in economic literature under that name. What 
Mtisehler and (»thers have identified as pcllucens could not be deter- 
mined without a genitalic examination of their specimens. Probably 
in many instances they ha<l mistaken argcntina for pclluccns. since 
bfdh species occur in the AVest Indies and Brazil. 

Aeeonling to Scott the favored host of pclluccns { — cistipcnnis) in 
I'uei-to Rico is the cowpea {Vigna unguirulata), and the species while 
iieqiient in lima Ix'ans seldom does serious damage. Potentially it is 

an insect of er-oiiMlnic Inipni | ;i|He. Tlie l;il\;ie are pliliiil iil\- tuxl 

borers but also bore into the stems and feed on the (lowers i^i their 
hosts. They attack. a>^ far as known, oidy Legumino>ae. 

I I Mil.l.l.A AIM.IM I\ \ K.Mir 
I'l.Mi ."., Kri.cuKs S Sf.- IM.AiK <i, Kl(iri!i:> IJ, 1.'! 

i'lDuh Ihi iiillin'(HM ZKr i> R (in p;irl, "v;ir. li"), Isis von < >ki'ii, vnl. |1, p. sCT, 1S4S; 

Iliinif Sof. Kilt. Hossicii(>, vol. ]<», p. 2^1, Wk. HI*, I'^^sl (new syimnyniy). 
FuuiliUn nifimtina Dyau, Inseeiifor Inscitinc .Menslnnis, vol. 7. i). I<>, VMW. 
J'lniihlhi lUfiiHiH I>YAi:, ln.'«eniinr Ifiseiline Menslnnis, vol. 7, ji. I", ini'.t ( new 

svimnyniv ). 



110 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

Male. — ^Anteniial shaft with even smaller black basal scale tuft than 
that of pellucens. Forewing gray without the reddish brown, inter- 
spersed scaling characteristic of typical examples of peUucens,' entire 
basal area to antemedian line dark fuscous gray (with but very slight 
dusting of whitish scales toward base in some specimens) ; this dark 
basal patch contrasted against the paler gray color of the remainder of 
the wing, extending from costa to inner margin and bordered out- 
wardly by a narrow whitish line. Otherwise not distinguishable, 
superficially, from pellucens. 

Alar expanse 15-20 mm. 

Genitalia (pi. 5, figs. 8-8c) without sclerotized subanal plate. Ter- 
minal projection of gnathos varying from round to pointed (pi. 5, 
fig. 8«) at apex. Harpe tapering to bluntly pointed apex; clasper a 
single, straight, slightly roughened, appressed spine, situated beyond 
middle of harpe. Aedeagus simple; cornutus a single, straight spine. 

Female. — Essentially like the male in color and markings except 
that the basal area of forewing is concolorous with or contrastingly 
paler than the remainder of the wing. A narrow dark line or a 
diffused dark shading outwardly bordering the obscure antemedian 
line. 

Alar expanse 15-23 nnn. 

Genitalia (pi. 6, figs. 12, 13) with signum well developed and con- 
sisting of a large pear-shaped cluster of thondike spines; sclerotized 
band behind genital opening, divided in the middle, simple (pi. 6, 
fig. 12) in Argentinian and Brazilian specimens, or armed with a pair 
of median, spinelike projections (pi. 6, fig. 13), rather long in West 
Indian specimens or short and disappearing in Mexican and Vene- 
zuelan specimens. 

Types. — In United States National Museum {argentina and 
eucasis). 

Type loeaUtics. — Tucuman, Argentina {argentina) ; Caracas, 
Venezuela {eucasis). 

Food plant. — Cassia spp. (reared examples in National Collection 
from Cassia hicapsularis and C. corymbosa). 

Distribution. — United States: Florida, Biscayne Bay (May), 
Coconut Grove (April); Texas, Brownsville (November). Mexico: 
Several examples reared from pods and blossoms of Cassia hicapsularis 
at Brownsville, Tex., quarantine station. Cuba: Baragua (March), 
Habana, Matanzas, Santiago Province. Puekto Rico: Bayamon 
(March, September), Vieques Island (April, July), Coamo Springs 
(April) , Aguirre Central (August) , San Gernnin (August), San Juan 
(November). Haiti: Petionville (June). Jamaica. Venezuela: 
El Valle (June). Brazil: Bahia (May). Argentina: Tucuman 
(March). 

Seventy-three specimens examined. 



1 1 f: ( . I- \ 1 



\i)ii.i.\ iii:i\i;i( II 



111 



In rolh'ctioiis [\\\> >|n'fU'> \\;\> ii[)[>VMVi\ liin>t i'l('(|iltiit 1 y lIIidiT \\\0 
luiiiH' fi) H III! lis. liotli (iri/) iitlna :nnl jtiJhiciits \\\\\v .iliiiiii tin- >iiiii<' 
tlisi i-iliiit'niii and ai't- ahnmlaiit in tlif \\'f-l Imlirs. (Ii niiili. t'loni ni;i- 
tfiial ai hand. /" Ihirt i\s si'cins (<» hi' rarer on tlic niiiinhimh 1 hn»u<_di- 
• Mit ii> riiiiiH' (irqeiit'iuii shows considcrMlili' variation in fcnialo 
ijt'nitalia. A\'c^i Indian -]ifiiin<Mi> haxc rather con-i lii iious spinidikc 
e.\ten>ions of the >eh'rot i/.i'd hand hi'hind tlie L^eiutal o|»enin<i. These 
are entirely laekiiii: in I>ra/,ilian specimens, and if one liad oidy those 
e\treine> he wonhl he jii-tilieil in a---nnii iii:" ihat they wcir at h'a>t 
I'aeially distinet. Ilowexi-r. \'i'ne/.nehiii and .Mexican e.\aiii|)lf- -how 
an intermedial*' form with veiv short |)roir(i ions, and Central Anieii- 
can >iiecimen>. when i'eco\ eicd in >nllicient nnnilicr-. will |Moltalily 
-liow all inter:iradation>. The male (.'enitalia ai-e remaiUahly nni form 
thronirhont the ranne of the specirs. exhihit injj; only minor individual 
\ariatioii- in the shape (d" the terminal projection of the ^nathos. 
The type (d' Dyar's t iira.sis i> oidy a -mall, -omewhat faded male (d" 
<U(jtiitin(i. 

FLNOKLI-A A<;AI'KI.I.A Schnus 
Pl.ATK <!. FKilliK 11 



/'ini'hilil <l<l>ll>< ll<l .•<( MAT >. /,cinln-ii;t. M <] . ."., Nu. L', \K J7. VJ'SA. 

Fi null, . — I*al|)i. liea(l. thorax, and foreuinLT whitish jiray: dark 
marking's drali ;:iay: (ran-\er>e antemeilian line (d' foicwinn white, 
detiiie(| chiefly liy it> narrow, dark outer hoi-der, sharjdy simiate, 
indent<'d a trifle just i»e|ow co-ta. more dccjilv at to]) of cidl and still 
more clecpjy at fol(| helow cell: »li-cal dot at end (d' c(dl oh-cur-e; 
white -uhterndnal line indented at Ncin (i and at suhmedian fold, hor- 
deied inwardly hy a di-tinct dark -hade a- hroad as the white line 
il-elf and outwardly hy a -iniilai'. fainter, dark shadinjf. the latter 
con-picnoii- only at apc\. Hind w in^^ ;,.. ji, th,. other species of 
h II 1x1 1 1 1 II. 

-Mar e\|.an-e j-j nun. 

(ienitalia (pi. Ci. lii:. 11) like tlio-e of intermediate example- of 
iiri/i nfiiiii ex<'ept that the sijjnmn i- ci .n-ider-ahl \ -lualler in propor- 
tion Id the -ize of the hursa. 

^ III"- — ^" I'niled Slates National Mir-euin. 

Tijin Jdriil'ilii. — '{'a^nis Cove, AlU'liiarle, (iala|»a;ros I.-land.s. 

I'linil j>liiiif . — I nknow II. 

Known only from the female type. SupeilicialK a ili-iinct -pecies. 
The female <i:enitalia. however, would indi<ate that <i<i,i i>, llii \~~ onlv a 
race of nrifintiiui. .\ male will he needi-d for exaci placement, and 
initil it i- axailahle we -hall have to treat nqii jn lln a- a -|iei-ies. 



112 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

FUNDELLA IGNOBILIS, new species 

Plate 5, Figures 9-9(/ ; Plate 6, Figure 14 

Male. — Antennal sliaft without any trace of black basal scale tuft. 
Otherwise partaking of the pattern markings of bath fcUucens and 
argentina; in some specimens dark basal patch of forewing round 
and reaching neither costa nor inner margin (as in typical pellucens) ^ 
in majority of specimens, however, basal patch occupying whole basal 
area (as in typical argentina) ; median and outer areas of wing averag- 
ing a trifle paler than in argentina and without the reddish-brown 
scaling of pelJucens. 

Alar expanse 13-20 mm. 

Genitalia (pi. 5, figs. 9-9<:Z) with gnathos terminating in a short, 
stout hook. Harpe with apex truncate; clasper moderately long, 
curved, and weakly haired at apex. Aedeagus with a single, long, 
strong, curved spine from below apex; cornutus a short, stout, curved 
thorn. 

Female. — Superficially similar to argentina except a trifle paler on 
the average. 

Alar expanse 15-22 mm. 

Genitalia (pi. 6, fig. 14) without spines adjacent to genital opening. 
Bursa copulatrix with signa, consisting of a pair of partially fused 
bands, each armed with a row of short, stout, thornlike spines; ductus 
bursae short and broad, with median area unsclerotized ; eighth seg- 
ment collar completely sclerotized except for a small, round, trans- 
parent spot on midventer, sclerotization extending to and over area 
behind genital opening. 

Type and paratypes. — U. S. N. M. No. 57185. Paratypes also in 
British Museum and in Cornell University and Janse collections. 

Type locality. — Oaxaca, Mexico. 

Food 2^1 ant. — Unknown. 

Described from the male type and 2 male and 4 female paratypes 
from the type locality; 4 male and 8 female paratypes from Tehua- 
can, Mexico (May, June, July) ; 3 male and 7 female paratypes from 
Orizaba, Mexico; 1 male and 6 female paratypes from Cordoba, 
Mexico (May); 1 female paratype from Guadalajara, Mexico; 1 
female paratype from Jalapa, Mexico ; 1 male paratype from Cayuga, 
Guatemala; 1 female from Costa Rica; 1 male and 1 female paratype 
from Santiago, Cuba (June) ; 1 male and 1 female paratype from 
Sierra Miestra, Cuba (May) ; 1 male paratype from Cuba without 
other locality label; 1 male paratype from Aguirre Central, Puerto 
Rico; and 1 female paratype from Petionville, Haiti. Most of the 
foregoing were in the National Collection under either pellucens or 
argentina. The species is quite distinct and easily recognized in 
either sex by its genitalia. 



THK (.K\IS II \|)|.l.l.\ IlKIMiK II 113 

FlM)i:i,I.A AlllMoltV D.Mit 
PiAiK I. Fi(.ri:i: r. : I'l ah: .'i, I"i<,i i;i:s in Id.-; I'l \ri:<;, FiuruE 16 

Fitmltlla iilu mora Dyau. I'mc. U. S. .\:it. Mus.. vnj. J7. p la'^. I'.il }. 

Mill*. — Aiit<'nna with siii:ill Klark -r.ilc lull al hasc of shaft. 
Forewiiiji with no or a \iry faini daik ha-al ])atch (when present 
coverinir i)asal area to anteiiieclian liiu') : aiitemcdian line whitish, 
veiT faint; subtenninal line white, faint but less oliscure than antc- 
niedian. without dark borders exeej^t for an inner and an oiiii r dark 
spot at inner margin of wing; veins from cell rather strongly outlined 
by dark scaling (the most conspicuous superficial character of the 
>pecies). A thick, dark (brownish) hair tuft covering outer surface 
of fore tibia ( ]•!. I. tig. '> » . a ihalc rliai-actcr not fnund in oilier species 
of the genus. 

Alar expanse 18-23 nun. 

Genitalia (pi. 5, figs. 10-lOr) with giuithos terminating in a broad 
tonguelike plate. Harpe somewhat tapering but with apex truncate; 
a stronii tuft of long- scales from costa: clasper long, curved, slender, 
with a few hairs at ajiex. Aedeagus with a paii- of long, curved, 
llattenecl spines from ai)ex; cornudis a long, straight, sh'uder >pinc. 

Fr/no/r. — Essentially like the male in color and markings. 

Alar expanse 18-23 nun. 

Genitalia (])\. Ct. fig. IC) with a pair of long, widely sjiaced, l)asally 
curved s]iin('s from sclerotiz('(l area iniiiiedialely beliind genital open- 
ini.'. r>iir-a id| Milat ri.\ with >igiia <<>ii^i^tiiig ol iwn ratliei' >hoi't 
baiid>. each arm«Ml with a row of long >pines. Ductus bursae bulged 
in the mi<Idle and with a strongly s( leiotized nie<lian collar. Collar 
of eighth segment partially sclerotizeil and fuscii veni rally. 

Ti//tr. — In United Stale- Xatioiial Mu~euni. 

T>/pc lornTtty. — Ori/.aba. Mexico. 

Food qyJaut. — UnkiHtw II. 

Ifistrihutioti. — Mi:.\ico: ()ri/,ai)a. .lalajia. 'I'eapa (December), Cor- 
doba (April. December). Cuernavaca (July), Gi atkm.m.a : Qiiiiigua 
iXfarch). Cayuga (January. May). Paiuiha (July). Costa Ivica: 
Juan Vinas (Xovember), 

•Nineteen sj)eciniens examined. 

Sup«'rficially the most easily di-i ingui>lied specie- in (he genus. 
The large forelibial tuft at once idetililies the male, and both sexes 
can be >e|iaraled bv the r.iiiiei- cori-|>icu<Hi- dark oiillining of the 
veins. The vein- .iie -iiiiilarly dark -c.iled in the other sjiecies, but 
(he contrast of the daik xcin- again-t the pale iiit er\ enular aieas is 
iiiofc marked in uln mnrd. 



EXPLANATION OF PLATES 

The drawings of figures 7-7c, 8-Sc, d-9d, 10-lOc, 11, 12, and 13 
for the plates accompanying this paper were made by Mrs. Eleanor 
A. Carlin, formerly with the Bureau of Entomology and Plant Quar- 
antine. Figures 1, 2, 3, 4, 5, 6-6c/, 14, 15-15f/, and 16 were drawn by 
Mrs. Sara H. DeBord, of the Bureau of Entomology and Plant 
Quarantine. 

Plate 4 

1-4, 6^7c, Fundella pellucens Zeller: 1, Side view of male head; 2, basal segments of male 
antenna, denuded to show depression in shaft; 3, same, showing scale tuft on 
shaft; 4, hind tibia of male; 6, wings of male showing venation; 6a, ventral 
view of anal pocket of male hind wing, opened to show scale tufts; 7, ventral 
view of male genitalia with aedeagus omitted; 7a, lateral view of tegumen, 
gnathos, subanal plate, and uncus; 7b, aedeagus; 7c, sternite and tergite of 
eighth abdominal segment of male. 
5, Fundella ahemora Dyar: Tufted foretibia of male. 

Plate S 

8-8c, Fundella argentina Dyar: 8, Ventral view of male genitalia with aedeagus omitted; 
8a, terminal projection of gnathos showing extreme of variation; 8^*, sternite 
and tergite of eighth abdominal segment of male; 8c, aedeagus. 
9-9d, Fundella ignobilis, new species: 9, Ventral view of male genitalia with aedeagus 
omitted; 9a, lateral view of tegumen, gnathos, and uncus; 9b, aedeagus; 9c, 
sternite and tergite of eighth abdominal segment of male; 9d, anellus. 
10-lOc, Fundella ahemora Dyar: 10, Ventral view of male genitalia with aedeagus omitted; 
10a, lateral view of tegumen, gnathos, and uncus; lOb, aedeagus; 10c, sternite 
and tergite of eighth abdominal segment of male. 

Plate 6 

11, Fundella agapella Schaus: Ventral view of female genitalia. 
12, 13, Fundella argentina Dyar: 12, Ventral view of female genitalia of type with bursa 
copulatrix omitted; 13, ventral view of female genitalia of West Indian speci- 
men. 
14, Fundella ignobilis, new species: Ventral view of female genitalia. 
15-15a, Fundella pellucens Zeller: 15, Ventral view of female genitalia; 15fl, collar of 
eighth abdominal segment of female, dorsal view. 
16, Fundella ahemora Dyar: Ventral view of female genitalia. . 
114 



U.S. GOVERNMENT PRINTING OFFICE: 1945 



U. S. NATIONAL MUSEUM 




6a 



.77NV 





PRCX:EEDINGS. vol. 96 PLATE 4 

-. 1 



MS 






ahe mora 



7 pellucens 




The Genus Fundella. 
For explanation sec pavrc 114. 



U. S. NATIONAL MUSEUM 



PROCEEDINGS. VOL. 96 PLATE 5 




ahemora. 



The Genus Fundella. 
For explanation see page 114. 



U. S. NATIONAL MUSEUM 



PROCEEDINGS. VOL. 96 PLATE6 




14 ifnobiliJ 



15 pellucens 16 ahemora. 



THE GENUS FUNDELLA 

For explanation sec pa^^:c 1 14. 



PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 




issued ft'-t^iVvA- sii»JB h the 



SMITHSONIAN INSTITimON 

U. S. NATIONAL MUSEUM 



Vol. % Wwhington: 1945 No. 3191 



A M:\V (iENUS AND TWO NEW SPECIES OF PEKCOID 
FISHES FROM XEW GUINEA, FAMILY CENTROPOMIDAE 



Hv Lion Aiv'i) P. Schultz 



Lt. Jmiik's R. Simon, U.S.N.R., irccntly donated to tho Unitod States 
National Museum a small collection of lishes jtriven to him hy his friend 
Capt. Rail)]) F. Honess, U.S.A.. who collected them in New Guinea 
in 1044. In this lot. lotalin*; '27 s])ecimens, ^Yere eijiht examples from 
fresh water that I consider to helonj; to an undcscribed pei-coid <;enus 
invoh in<_' two new species. These are described herein. The other 19 
.sp»'cinicn< arc of known foiins oc('nn'in;Lr in the re<ri<>ii. 

XENAMBASSIS, new genus 

Geiwtypi . — Xr)u//)iha.y.s/',<< lifrytpss'/. new species. 

Body ronij)ressed. covered with cveioid scales of moderate si/e every- 
where except on top of head: basal half of caudal fin scaled and a few 
-cales on base of peel oi'al : lateral line arched ^\\^'\• j)e<'loral fin so that 
It is concurrent with doisal pi'ofile. then extendin;:; alon^ midaxis of 
body posteriorly; each lateral line tube straiefht. not extendine; entire 
len^jth of scale ; anterior profile concave ovci- orbits : intei-orbital space 
a little convex; a low predorsal ridejc exteiidinir alon^r middorsal line 
endin«r at occiput: mouth teiminal. a little obruiue; premaxillary 
diLditly [irotractile : maxillaiy withoiil supplemental bone, mostly t'X- 
posed, o(dy the doj-sal edp- slipi)in;j: sli^dit ly mulei- ed<fe of preorbital 
bone; maxillaiy with its {posterior ed^'e conca\e: jaws e(|ual or nearly 
-CI. the lower sli^htlv in front id' upper: iccth in both jaws in a \illi- 
t'orm band, with the out«*r row consisting; of somewhat e;darjie(l conical 
teeth, the band of teeth becoming' narrower on sides of jaw. endinjj 
in onlv one or t wo rows on lower jaw : vomer with a patch of villi form 
teeth and palatines with a nairow row of \ illifoini teeth : no teeth on 

620241— 4.^ 11'' 



116 PROCEEDINGS OF THE NATIONAL MUSEUM vol.98 

tongue; cheek and operculum scaled; posterior and lower margins 
of orbit denticulate and two denticulate ridges on preorbital bone 
above maxillary, the upper one forming part of the serrae around 
orbit ; preoperculum with double edge, the outer one serrated on both 
posterior and ventral borders, the inner one serrated at lower angle 
and a little on each side of the angle above and forward; inter- 
operculum denticulate ; operculum not spinate ; gill membranes united 
far forward, free from isthmus; branchiostegals 6; pseudobranchiae 
present ; gill rakers of moderate length, of rather heavy build ; a single 
dorsal fin with the spiny portion scarcely longer than soft portion of 
VIIT or IX spines, the second or third longest, the next to the last 
shortest; spinous dorsal fin preceded by a short recumbent spine, 
directed forward, hidden in the skin beneath the scales ; anal with 3 
spines and usually 10 soft rays ; both fins with a sheath of scales along 
their bases, one scale in width along dorsal and one or two along anal, 
these fins partly depressible between the sheath ; pectorals ii, 13, asym- 
metrical, upper 2 or 3 branched rays longest ; pelvic fins, I, 5, inserted 
under base of pectorals and a little in front of dorsal origin; pelvic 
spine slender, not reaching to anus and about two-thirds length of 
first branched pelvic ray, which ends in a short filament; axillary 
scale of pelvic small, about length of diameter of pupil; caudal fin 
forked, lobes more or less pointed. 

Other characters are those of the genotype, X. honessi, described 
below. 

This new genus is related to the percoid fishes usually referred to 
the family Centropomidae but sometimes separated from them and 
grouped in the family Ambassidae or Chandidae. Xenainbassis is 
especially close to Tetra^entrmn Macleay, Synecliopterus Norman, 
and Amhassis Cuvier and Valenciennes, differing from them as in- 
dicated in the accompanying key. Ambassis differs from the new 
genus and from Tetracentrum and Synechopterus by having the first 
dorsal fin made up of VII spines and joined to the base of the first 
spine of the second dorsal, whereas in the other three genera the first 
dorsal is joined at least halfway out the last spine so that the two 
fins are continuous. Chanda Buchanan-Hamilton, 1822, with G. lala 
as the type as restricted by Fowler (Proc. Acad. Nat. Sci. Philadel- 
phia, 1905, p. 500) differs from the above-mentioned genera chiefly in 
having 14 to 17 soft rays in the anal fin. Pseudoarribassis Castelnau, 
1878 {— Austrochanda Whitley, 1935), a substitute name supposed by 
Whitley to be preoccupied by Pseudarribassis Bleeker, 1876; and Vel- 
anihassis, proj)osed by Whitley (Rec. South Australian Mus., vol. 5, 
No. 3, pp. 356-365, 1935) , are here considered as subgenera of Amhassis 
Cuvier and Valenciennes. Indeed, they may even be synonyms of 
Ambassis, since the generic differences appear so slight. I also refer 



NEW PERCOID FISHES I'HOM NEW GUINEA SCHULTZ 117 

Acanthoperca Castoliuiu, 1S78, ami lilandoirskieUa Irediile and Whit- 
ley, 1932, as subgenera of Aiiibassh. Priopidichthys Whitley, 1935, 
with teeth on the tongut.'. may well he a valid grnus, as none of the 
other irenera seem to have lingual teeth. 

Named Xena//>h(issi.s, meaning a strange or different Ambassis. 

KEY TO THE GENERA AND SPECIES OF NEW GUINEA 
CENTROPOMIDAE RELATED TO TETRACENTRUM 

la. Anal rays IV, f>; dorsal rays IX, 10, dorsal tiu loiitiiuious ; preorbital witli 2 
serrated ridges; suborbital and postorbitnl ring of bones witb denticula- 
tions ; inner double edge of preoi>ercular bone strongly toothed at angle 
and along its lower edge; outer edge of preoperculum strongly denticulate 
aluug its entire border; least depth of caudal peduncle Ir.i in its lenytii : 
profde concave over orbits ; pelvics inserted under i^)ectoral base; postorbital 
length of head 1.1 in length of caudal peduncle. 

Tetracentruni ' apogonoides (Macleay) 
lb. Anal rays 111, ."S lu 11 (see table 1). 
2a. Dorsal fin continuous, no deep notch in front of last spine, membrane be- 
twt^n la.st two spines connected over halfway out last dorsal spine; sec- 
<»nd dorsal spine not reaching anywhere near base of last dorsal spine 
when fin is dtpressed. 
3n. Orbital rim without serrae; profile over orbits convex; inner ridge ot 
preoperculum with 1 or 2 serrae at angle; pelvics inserted just behind 
pectoral base; anal origin under next to last dorsal spine; least deptli 
of caudal peduncle 1% in its length; postorbital length of head 1.3 in 
length of caudal peduncle; dorsal rays IX, 10; 14 gill rakers on lower 
half of first gill arch ; pectt)ral with 14 rays. 

Synechopterus caudovittatus Norman ^ 

:ib. Orbital rim denticulate; profile over orbits a little concave; inner ridge 

of preoperciilinn with lower and posterior sides near angle denticulate; 

Itelvic iiLsertion under pectoral fin base; anal origin under base of last 

dorsal spine or base of first soft dorsal ray. 

.}a. Dor.sal rays VIII, 10 or 11; a blackish band along nudaxis of body 

commencing behind head and becoming more intense on caudal 

[jeduncle, its width about equal to that of pupil, ending in a dark 

blotch at base of caudal fin Xenanibassis honessi, new speci<>s 

■'ih. Dor.sal rays IX, 10; midaxis of body with a narrow dark streali on 
caudal region not ending in a large dark lilol( h at base of caudal tin. 

Xenanibassis simoni, new specie's 
2h. Dorsal lln dcply notclied, first portion connected at ba.se of first spine of 
second dorsal fin; dorsal rays VII-I, 8 to 14; second dorsal spine long, 
slender, when depressed Its tip reacliing past base of last dorsal spine. 

Aiubassis ' Cuvier and Valenciennes 



' Whitlfy, 1935, propoHed VcgamhaHsit to replace Tctracentrum, which ho Hnid was pre- 
oi('ii|il.-.| l)y Trtrnrrntron BrainT, 18t!.1. but the apt-llinR U not In conflict, acconllnK to 
oplnioiiH 147 anci \\H of the Intcriuitioiial CotnmlHRlon on Zoological Nomenclature, and bo 
Tctracentrum muHt Htand aH a valid name. 

' Copela. lOa.',. No. 2. pi>. 01-0:j. flj;. 1. 

•For a key to the Hpecles of AmhatitiH Hee W'elxT and (!<• Iti-aufort, FIhIh-h of the ludn 
.\ufltrallnn Arrhlpelajco, vol. 5, p. 398, 1029. 



118 



PROCEEDINGS OF THE NATIONAL MUSEUM 
XENAMBASSIS HONESSI, new species 

Figure 3 



HoJoti/pe.—V.S.^M. No. 122830, a specimen 87.6 mm. in standiird 
length, collected in 1944 by Capt. Ralph F. Honess, U.S.A., in either 
the Samboga or the Giriia River at Buna, New Guinea (long. 148° 
30' E.; lat. 8°45' S.). 

Paratypes.—U.S.^M. No. 122831, 3 specimens, T7.5, 80.5, and 82 
mm., collected along with the holotype and bearing same data; 
U.S.N.M. No. 121832, 2 specimens, 58 and 87.5 mm., bearing same 
data as holotype. 

Descriptio7i. — Detailed measurements were made, and these data, 
expressed in hundredths of the standard length, are recorded first for 
the holotype and then for two paratypes in parentheses, respectively. 
Standard lengths in millimeters 87.6 (82; 87.5). 

Table 1. — Counts recorded for certain species of Centropomidae from New Guinea 





Number of fin rays 


Number of scales 




Sppcios 


Dorsal 


Anal 


Pectoral 


Lateral line 


Above 

lateral 

line 


Below 

lateral 

line 




VIII, 

10 


VIII, 

n 


IX, 10 


III, 10 


IV, 9 


ii, 12 


ii, 13 


30 


31 


32 


4 


7 


8 








1 
1 

2 


1 
6 
2 


1 






1 

3 

1 






1 
1 
6 
2 


1 
1 
5 
1 




Syn echopter us cau dorittaius 






1 


11 
4 


3 

1 


1 






1 


5 


1 


XeTianibassis siiuoni 






1 















Greatest depth of body 46.2 (45.1; 44.6) ; length of head 37.7 (37.8; 
37.4) ; length of snout 9.25 (9.39; 10.2) ; diameter of eye 11.0 (11.3; 
11.8) ; least width of interorbital space 9.82 (9.76; 9.02) ; length from 
tip of snout to rear tip of maxillary 13.7 (13.4 ; 14.7) ; postorbital length 
of head 18.3 (18.8; 17.7); least width of preorbital opposite tip of 
maxillary 2.28 (2.20; 2.28) ; least depth of caudal peduncle 14.4 (14.4; 
13.7) ; length of caudal peduncle from base of last anal ray to mid- 
caudal fin base 18-8 (18.7; 17.4) ; length of procumbent embedded spine 
at origin of dorsal fin 4.56 (4.27; 4.34); lengih of longest ray of 
pectoral fin 25.5 (27.1 ; 25.7) ; longest soft ray of pelvic fin to end of fila- 
ment 25.7 (26.8; 25.1) ; length of pelvic spine 15.6 (16.5; 16.6) ; length 
of first dorsal spine 8.10 (6.95; 7.77) ; of second dorsal spine 19.6 (19.8; 
18.3) ; of last dorsal spine 13.8 (14.5; 15.2) and next to last spine 13.5 
(13.0; 13.2) ; longest soft ray of dorsal fin 20.5 (20.1 ; 21.7) and of anal 
fin 21.6 (22.0 ; — ) ; length of first anal spine 8.56 (6.22 ; 8.34) , of second 
12.0 (12.8; 15.6), and of third anal spine 14.8 (14.0; 14.8); longest 
caudal fin ray 34.2 (33.5; 30.0) ; shortest midcaudal fin ray 16.8 (17.4; 
16.1) ; length of longest gill raker on first gill arch 3.77 (4.27; 3.65) ; 



NEW PEHCOll) FISHES IKOM NEW CTINEA — StlllLrZ 



IIU 



(listaiK-e from ti|) of snout to dorsal orijriii 4(').S (47.0; 47.4) : snout to 
niuil oii<:iu G")!) (G4..'5; 65.7): snout to pectoral insertion 34.2 (30.8: 
3(5.0); snout to jjelvie insertion 3i>.0 (39.0: 11.5): snout to center of 
:inus .'m.C) (5r..7 : ."rj.l ) : ceiitei- of anus to anal ori<i;in 7.<t.") (7.32: ,s.()() i. 

The follo\vin«r counts were made, respectively : Dorsal rays VI 11. 11 
(VIII. 10: VIII. 11: VIII. II: \TI1. 11: VIII. 11) : anal rays III, 10 
in all six specimens; pectoral rays ii. 13 in each side of all six s[)eci- 
mens: pelvics always I. 5: iriH rakers 6+1 + 13 in all the types; scales 
in lateral line 31 (30; 31; 31: 30; 30) ; scales ahove lateral line 4 (4:4; 
4; 4; 4) and helow lateral line to anal ori«;in 7 (7: 8; 7: 7; 7) ; zigzag 
.scales around caudal i)eduncle 15 (l6: IC); 16; 15; 16) ; branched rays 
of caudal fin always 8 + 7. totaling i:». 



\^fl^_v 




^^ 



Figure 3. — Xenambassis honessi, new species: Holotype (U.S.N.M. No. 122830), standard 
length X7.6 mm. Drawn by Mrs. Aimc \I. .\wl. 

Depth of body about 2'/). head 2%, both in standard length; snout 
shorter than eye 3»'- to 4. eye 3yi„ to IP/-,, both in length of head; in- 
terorbital about e<|ual to snout; maxillary with its rear margin a little 
concave, not quite reaching to below middle of pupil; three series of 
scales on the che<.»k ; dorsal origin over rear of base of pectoral Hn ; 
first dorsal spine short, second aixl third of nt-arly e(|ual length, those 
following gradually shorter, the next to last sjjine shortest, almul 
three-fourths length of last : anterioi- soft rays of both dorsal and anal 
tins longest, longer than any of the spines in the same fin>; first anal 
>pine about two-thiids length <»f second and third, the ladcr s|)ine 
usually a trifle longer than the second ; -pines of dorsal fin about ecjual 
to f)ostorbital length of head: pelvic spine a little shorter than |)o>t- 
orbital length of head and the first soft ray a little pr<i(|uccd. reaching 
about opjiosite the anal origin: pectoral fin iieaily or (|nitc reaching 
lo opposite anal origin; vertebrae 10 i 17. 



120 l'lU)C;iOEDlNCiS OF THE NATIONAL MUSEUM voi-. »b 

OoTonrJion. — Goncrnlly brownish, with the center of each scale 
pnU'i-; a pule, hroiul, wedge-shaped bh)(ch extends from in front of 
anus to pelvic base, upward to behind pectoral fin base; a blackish 
laleral band extends alono- niidaxis of body, more intensely blackish 
l)os(eriorly, eniliiig in a black blotch at base of caudal lin ; a dark blotch 
about size of eye or smaller occurs just dorsally to anterior base of 
anal fin ; all (ins dusky, the pip;ment more intense basally on interradial 
membranes ol" soft I'ays of each median lin; clieek pale; operculum 
with dusky blotch above and below, w hich are palei- areas; peritoneum 
black. 

lieinarka. — This species may be separated from all rehited forms by 
the foregoing key. 

Named honessl in honor of its collector, Capt. Ilalj)h F. Honess, 
U.S.A. 

XENAMBASSIS SIMONI. ni-w KpetiiN 

Figure 4 

llolotype. — U. S. N. M. No. l'J282.S, a male specimen 80 nnn. in 
standard length, collected in either tlie Samboga or the Girua Kiver 
at Huna (long. 148^'U)' E., lat. 8°45' S.), New Guinea, by Capt. 
Uali)h V. lioness, U.S.A., in 1944. 

Pamti/pe. — U. S. N. M. No. 122829, a fenuile si)ecimen, 08.5 nmi. 
in standai'd length, taken along with the holotype and bearing 
same data. 

Description. — Detaiknl measnrements were made, and tliese data, 
expressed in hundredths of the standard length, are recorded first 
for the holotype and then for the para(yi)e. Standard lengths in 
millimeters 80 and (58.5. 

(JreatesI depth of body 45.0 and 48.8; length of head ;H).9 and 40.7; 
length of snout 9.25 and 9.78; diameter of . eye 10.9 and 13.0; least 
width of inlerorbital space 9.38 and 10.4; length from tip of snout 
to rear tij) of maxillary 13.5 and 14.2; post orbital length of head 
17.4 and 19.9; least width of i)reorbital opposite tip of maxillary 
2.00 and 2.34; least depth of caudal i)e(luncle 15.1 and 15.0; k'ugth 
of caudal peduncle from base of last anal ray to midcaudal tin base 

17.3 and 18.7; length of embedded procumbent dorsal spine at dorsal 
origin 4.25 and 3.94; length of longest ray of i)ectoral fin 25.3 and 
24.8; longest soft ray of pelvic lin to end of lilament 27.1 and 2G.7; 
length of pelvic spine H').2 and 17.4; length of first dorsal spine 8.50 
and 7.44; of second doi-sal spine 19.2 and 21.3; of last dorsal spine 

14.4 and 11. 5; of next lo last dorsal spine 14.1 and 11.5; longest soft 
ray of dorsal fin 19.5 and 19.0; of anal lin 20.0 and 20.3; length of 
first anal spine R.50 and 7.88; of second 13.3 and 14.9; of third anal 
spine M.4 and 14. 0; longest caudal lin ray 33.1 and 34.0; shortest 
midcaudal lin ray 10.2 and 10.3; length of longest gill raker on first 



NEW PERCOin FISHES FROM NEW (UTIXEA SCHULTZ 



121 



g\\\ arch ;i.l3 iind i5.65; (.listanoe from snout tip to dorsal ori«xin 44.4 
and 49.G; snout to anal origin 64.0 and 04.8; snout to pectoral inser- 
tion 33.8 and 37.7; snout to pelvic insertion 39.3 and 43.0; snout to 
center of anus 56.8 and 58.8; center of anus to niial orif^iii 7.02 
and 5.55. 

The following counts were made, respectively: Dorsal rays IX, 10 
and IX, 10; anal rays III, 10 and III, 10; pectoral rays ii, 13-ii, 13 
and ii, 13-ii, 13; pelvics always I, 5; gill rakers on first arch 6+1 + 13 
and 6 + 1 + 13; scales 30 and 31; scales above lateral line 4 and 4, and 
below lateral lino T and 8; 7,igzag scales around caudal peduncle 
15 and 10; branched caudal fin rays 8 + 7 and 8 + 7. 






Fu;lre 4. — Xenambassis simoni, new species: Holotype (U.S.N. M. No. 122828), standard 
length 80 mm. Drawn by Mrs. Airae M. Awl. 

The .shape of the body, length of fins, and otiior characteristics 
except certain counts and coloration are so similai- to X. hoiusK} 
that it is not deemed necessary to repeat thom here, since dct.iili'fj 
measurements are given ai)ove. 

Coloration. — (leneral coloration light brownish with centers of 
.scales paler; lateral line i)lackish on caudal region but no black 
blotch at base of caudal fin; area behind pectoral ami above |)elvic 
fins pale to midaxis of l)0(ly; median fins dusky, more intensely 
pigmented on interradial membranes basally; paireil fins light dusky, 
pelvics darker distally; peritoneum black. 

Rrwarh-H. — This new species may be distinguished IVoiu <losely 
ulatefl forms by the foregoing key. 

Named nlwoni in honor of Lt. James R. Simon, U.S.N.H., formerly 
of the Wyoming fiame and Fish Commi.ssion, who donated the 
specimens collected by Captain Honess to the National Museum. 

n » «OVIItllMCNT PXINTINa OfrirC l>4> 



PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 



issued |^r»(vA- QlJfil h ^' 




SMITHSONIAN INSTITUTION 
U. S. NATIONAL MUSEUM 

Vol. 96 Washington: 1945 No. 319 2 



THREE NEW SCIAEXID FISHES OF THE GENUS OPHIO- 
SCTON FROM THE ATLANTIC COASTS OF CENTRAL 
AND SOUTH AMERICA 



By Leonard P. Schultz 



During my studies of the sciaenid fishes of Venezuela I found a 
specimen of phioscion (U.S.N.M. No. 86710) from Uruguay, col- 
lected by the late Dr. Hugh M. Smith in 1922, with 28 soft rays in the 
dorsal fin. This count did not agree with the statements of recent 
authors who have reported on fishes from Panama and from the 
West Indies under the name of adustus. Although Jenyns and 
Agassiz each counted 28 soft dorsal rays for adustus from Maldonado 
and Montevideo, Jordan and Eigenmann in their review of the 
Soiaenidae (Rop. U. S. Comm. Fish and Fisheries for 1886, pt. 14, 
p. 403, 1889) decided that Agassiz's count was incorrect and should 
have been 22 or 23, or perhaps 18 or 19. Thus the species of the western 
Atlantic were confused in the first review and the name adustu,'i has 
been applied to two or three Atlantic species. 

This contribution discusses the relationships of the western Atlantic 
species of the genus Ophioscion and describes three new species from 
Panama, Venezuela, and Brazil. 

Drawings of figures 6 to 8 were made by Mrs. A. M. Awl. 

GenuH OPHIOSCION Gill 

OlihioMfion (JiLr.. I'roo. Afnd. Nat.'Sci. I'liiliidcliihia, vol. ITi, p. 1(V5, 1863. (Type: 
OphioJtritm typU-HH GUI, Itasj'd on U.S.N.M. No. iJ2SGl, west con.st of Panama.) 

No at(eni})t is made to include heroin the numerous refenMiccs to 
jocjilitv r*"'>!-.U of ~t,ifi<-t; nf (li(> gi'iuH f) pit i/>sc'n>ti from the western 

123 
62o4-12 -45 



124 PROCEEDINGS OF THE NATIONAL MUSEUM vol. b6 

Atlantic Ocean, because this would require a study of the specimens 
on which such reports were based, and these specimens are not now 
available. 

The genus Ophioscion is closely related to Bairdiella Gill and 
Stellifer Oken. Meek and Hildebrand (Marine fishes of Panama, 
vol. 2, p. 611, 1925) separated Stellifer from Bairdiella and O'phioscion 
in their key on the basis of the skull being "excessively cavernous, 
spongy to the touch." I had some difficulty separating the various 
species referred to these genera by this character, and so 1 made a 
dissection of the upper surfaces of the skull, removing the scales and 
skin, thus exposing the nature of the cavernous skull. Bairdiella 
chrysura (Lacepede), Ophioscion typicus Gill, and Stellifer rastrifer 
(Jordan) all have cavernous skulls dorsally, and also around the orbits 
occur narrow bony bridges or stays supporting the overlying skin 
and scales. Stellifer has a broader interorbital space, and thus the 
caverns are a little broader and by touch can be felt a trifle more easily 
than the slightly narrower caverns in the other two genera. My dis- 
sections indicate that the caverns are weJl developed in all three gen- 
era ^ and are of little value as a diagnostic character in the separation 
of these three genera, especially if the specimens are weU hardened 
in preservation. 

Ophioscion is said to differ from Bairdiella by having the lower 
spine of the preopercle pointing straight backward and a little down- 
ward, whereas in Bairdiella it is said to be hooked downward. The 
lower preopercular spine in these genera is so variable among the 
various species of Bairdiella and Ophioscion that I cast serious doubt 
on its usefulness as a character. None of the species of Ophioscion has 
any of the preopercular spines hooked downward. However, some 
individuals, especially young examples of Bairdiella chrysura^ like- 
wise do not have the lower preopercular spine hooked downward, 
although in adults of Bairdiella that spine is hooked downward. 

I have searched for characters to separate these three genera but 
have found indications of overlapping, and so the following char- 
acters are not wholly satisfactory, although the genera can be sepa- 
rated by them when taken together : 

Bairdiell'G, with an obliquely terminal mouth; both jaws of nearly 
same length; lower jaw with minute teeth in a narrow band of two or 
three rows forward and in a single row of slightly enlarged teeth 
posteriorly ; the pair of small pores at tip of chin close together and 
lying more or less in a shallow depression (fig. 5, &) and the margin of 
the snout lacking the small lobes at each side of the median pore 



* I removed the skin and scales from specimens representing various species usually 
referred to the following genera and found the dorsal part of the skull to be cavernous : 
TJmhrina Cuvier ; ilicropogon Cuvier and Valenciennes ; Plagioscton Gill ; Macrodon Schinz ; 
Gynoscion Gill ; Corvula Jordan and Bigenmann ; Larimus Cuvier and Valenciennes. In 
Mentlcirrhus Gill the caverns were much smaller than in the other genera examined. 



THREE NEW SCIAEXID FISHES — SCHULTZ 



125 



(fig. 5, a) ; the «2:ill rakers moderately long and slender, contained less 
than twice in the diameter of the eye; the first soft ray of the pelvics 
ending in a filament. 

Stellifer^ with an oblique mouth, somewhat intermediate between 
BairdieUa and Ophios^cion^ a little more inferior in position than in 
BairdieUa but not ventral in position as in Ophioscion; the snout 
projecting a very little in front of the tip of the lower jaw; low^er jaw 
with teeth in a narrow villiform band, the inner row of which is a 
little enlarged; the pair of small pores near the tip of chin separated 
by a small bony knol) (fig. o, d) ; the margin of the snout lacking a 
lobe at each side of the anterior median pore near margin of snout 









b d 

Figure 5. — Diagrammatic sketches of the tip of the snout and of the anterior part of the 
underside of the lower jaw of three species of sciaenid fishes: a. Snout tip of BairdieUa 
chrysura (Laccpede); h, lower jaw of B. chrysura; c, snout tip of Stdlifer rastrifer (Jor- 
dan); d, lower jaw of S. rastrifer; e, snout tip of Ophioscion typicus Gill (type, U.S.N.M. 
No. 22861); /, lower jaw of 0. typicus. 

(fig, 5, c) ; the gill rakers slender and long, equal to eye or contained 
in it fewer than two times; the first soft ray of the pelvic fins ending 
in a filament. 

Op/i/osc!on, wilh the mouth in a somewhat ventral position, the 
lower jaw included and jaws nearly horizonlal, with the snout pro- 
jecting a little beyond the tip of the lower jaw ; lower jaw with a wide 
Ijand of villiform teeth, none of whicii is enlarged; the pair of small 
l)ores at midtij) of hnver jaw close together and usually lying in a 
sliallow depression (fig. 5, /) ; the front margin of the snout bearing 
a short, blunt lobe each side of the anterior median |)()re (fig. 5, e) ; 
file gill rakers usually short, not slender, and coiitaiiied more than 
•J. 4 to times in the eye; (he first soft ray of the |»el\ir liiis ending in 
a filament, usually white. 

Thorugh the courtesy of Dr. Tiiomas Barbour, of the Museum of 
Comj^arative Zoology, I ha\e been able to examine some of (lu^ speci- 
mens dewribed by Jordan and Kigenmann in l.ss!) in llieii- review of 



126 PROCEEDINGS OF THE NATIONAL MUSEUM vol.98 

the Sciaenidae. ^''Stellifertts naso Jordan," based on specimens from 
Cachiura, Brazil (M.C.Z. Nos. 4583 and 10808), definitely belongs 
to the genus Ophioscion. Specimens of Stellifer microps Steindachner 
(M.C.Z. Nos. 4581 and 1031), from Para and Tonteboa, are Ophioscion 
microps (Steindachner). 

Jordan and Eigenmann identified M.C.Z. No. 22417 as Sciaena 
adusta Agassiz, but my reexamination of this specimen indicates that 
it is Ophioscion punctatissimus Meek and Hildebrand. 

The following key is intended to separate the species of Ophioscion 
occurring in the Atlantic along the coasts of Central and South America 
and in the West Indies : 

la. Dorsal rays usually X-I, 28 or 29; anal rays II, 8; gill rakers on first gill 
arch 9+1+16; scales 50 to 57; dark streaks commencing at upper part of 
back, passing forward and obliquely downward, then a little above lateral 
line bending abruptly downward, almost vertically, disappearing near mid- 
axis of body ; tip of spiny dorsal fin dark, base of dorsal fin with a narrow 
pale or whitish band, above which the fin is abruptly darker ; opercle dusky 

(mouth of Rio de La Plata) Ophioscion adustus (Agassiz) 

16. Dorsal soft rays fewer than 25. 
2a. Anal rays II, 9. 

3a. Dorsal rays X-I, 21 ; gill rakers on first gill arch 8 or 9+1+13, totaling 
22 or 23 ; scales about 45 or 46 ; eye diameter 1.2 to 1.8 in interorbital 

space Ophioscion brasiliensis, new species 

8&. Dorsal rays XI-I, 21 or 22; gill rakers on first gill arch 9 to 11+1+17 to 
19, totaling 28 to 31 ; scales about 47 to 51, eye diameter from 1% to 2% 

times in interorbital space Ophioscion microps (Steindachner) 

26. Anal rays II, 7 or 8. 

4a. Scale rows above lateral line usually 51 to 57. 

5a. Gill rakers on lower part of first gill arch usually 16 to 18 including 
rudiments ; dorsal rays XI-I (rarely XII-I) 21 or 22 ; anal rays II, 8 ; 
scale rows 52 to 54; color more or less plain grayish above, paler 

below Ophioscion venezuelae, new species 

56. Gill rakers on lower part of first gill arch 11 to 13 (see table for counts) . 
6a. Dorsal rays X-I (occasionally XI-I) , 22 to 24 ; anal rays II, 7 (rarely 
II, 6) ; gill rakers 7 or 8+1+11 to 13; scale rows above lateral 
line about 54 to 57. 

Ophioscion punctatissimus Meek and Hildebrand 
66. Dorsal rays X-I, 20 or 21 ; anal rays II, 7 (rarely II, 8) ; gill rakers 
7 to 9+1+13 ; scale rows above lateral line about 51 or 52. 

Ophioscion panamensis, new species 

46. Scale rows above lateral line 45 to 49 ; anal rays II, 8 ; dorsal rays XI-I, 

21 ; gill rakers 8 or 9+1+14 or 15; scales 46 to 49; eye diameter 1.0 to 

1.1 in interorbital space Ophioscion naso (Jordan) 

OPHIOSCION ADUSTUS (Agassiz) 

Corvina adusta Agassiz, in Spix and Agassiz, Selecta genera et species . . . 

Brasiliam . . ., p. 126, 1831 (Atlantic Ocean off Brazil).— Jenyns, The 

zoology of the voyage of H. M. S. Beagle, pt. 4, Fishes, p. 42, 1842 (Maldonado 

and Montevideo). 
Ophioscion adustus Tortonese, Boll. Mus. Zool. Anat. Comp. Univ. Torino, ser. 3, 

vol. 47, No. 100, p. 130, 1939 (Rio de Janeiro and Montevideo). 



THREE NEW SCIAENID FISHES — SCHULTZ 



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128 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

? Ophioscion icoodwarcU Fowled?, Proc. Acad. Nat. Sci. Philadelphia, vol. 89, 
p. 311, fig., 1937 (Port-au-Prince, Haiti). 

Corvina adusta Spix, pi. 70 in Spix and Agassiz (1831), has but 
X-I, 19 dorsal fin rays and 52 vertical scale rows above the lateral 
line; the anal rays as shown are II, 7. Obviously Agassiz did not 
describe the fish figured by Spix, since he gave dorsal rays as X-I, 
28 and anal rays II, 9. The most significant character in certain re- 
spects as clearly shown in plate 70 is the direction of the scale rows 
below the lateral line. None are shown parallel with the axis of the 
body, but the scale rows run obliquely upward and backward to the 
lateral line and the scale row arising from the rear base of anal fin 
meets the lateral line just behind a vertical from the rear base of the 
dorsal fin. The direction of these scale rows strongly suggests that 
plate 70 in Spix and Agassiz (1831) may be some species of 
Plagioscion^ perhaps near P. pauciradiatus Steindachner (1917). 

The following references list O. adustus^ but their counts disagree 
with those for the true adustus: Ribeiro, "Fauna Brasiliense Peixes," 
Arch. Mus. Nac. Rio de Janeiro, vol. 17, family Sciaenidae, p. 23, 1915, 
and.Devincenzi, Ann. Mus. Nac. Montevideo, ser. 2, pt. 5, p. 239, 1924. 
It must be concluded, therefore, that their descriptions must apply 
to some other species or that they were erroneously drawn up. 

The description by Jenyns fits very well the specimen before me 
(U.S.N.M. No. 86710), which is 120 mm, in standard length and was 
collected by Dr. H. M. Smith in Uruguay in 1922. 

Berg (Ann. Mus. Nac. Buenos Aires, vol. 4, p. 52, 1895) listed 
Sciaena adusta (Agassiz), but his counts did not agree with those of 
Agassiz or of my specimen, and they need reexamination to determine 
the identity of his material. 

Ophioscion tuoodtcardi Fowler {loc. cit.), described from Haiti in 
1937, probably is a synonym of adustus. I have not seen Fowler's 
types, but the number of fin rays places it with adustus, and the white 
area along base of dorsal fins abruptly set ofi' by the blackish area 
distally along basal part of dorsal fin rays, as in our specimen 
(U.S.N.M. No, 86710) from Uruguay, indicates that woodwardi and 
adustus are the same. 

OPHIOSCION BKASILIENSIS, new species 
FiGUEE 6 

Uolotype. — U.SN.M. No. 87742, one specimen, 77 mm. in standard 
length, taken over a sand bar at Santos, Brazil, September 12, 1925, 
by Dr. Waldo L. Schmitt. 

Paratype. — U.S.N.M. No. 122611, one specimen, 89 mm., taken with 
the type and bearing same data. 



THREE NEW SCI.VENID FISHES — SCHULTZ 129 

Descriptioji. — Certain meapurcmeiits were made, and these data, 
recorded below, are expressed in hundredths of the s(an(hutl k'ngth, 
fii-st for the holotyjie, then for the paratype in parentheses, respec- 
tively. Standard len<rths in millimeters, 77 ^89). 

Length of head 33.8 (33.2); greatest depth of body 32.5 (30.8); 
diameter of eye 7.53 (7.42) ; length of snout 9.22 (8.70) ; distance 
from front of upper lip to rear tip of maxillary 12.2 (11.8); least 
preorbital width 4.1G (3.93) ; postorbital length of head 18.8 (18.4) ; 
width of bony interorbital space 9.74 (9.10) ; length of caudal peduncle 
or distance from base of last anal ray to midcaudal fin base 22.1 (23.G) ; 
least depth of caudal peduncle 10.0 (9.r>5) ; length of base of second 




Figure 6. — Ophioscion brasiliensis, new species: Ilolotype (U.S.N.M. No. 87742). 

doi-sal fin 32.5 (31.2) and of base of anal fin 12.5 (12.9) ; longest dorsal 
spine 175 ( — ) ; length of second dorsal spine 12.3 (11.8); length of 
second anal spine 16.2 (13.8) ; longest ray of pectoral fin 24.4 (22.5) ; 
longest soft ray of pelvic fin 25.1 (25.8) ; length of pelvic spine 12.1 
(11.7); longest midcaudal fin ray 2G.0 (23.0); length of longest gill 
raker 3.64 (2.81) ; distance from snout tip to dorsal origin 37.1 (36.5) 
and to anal origin 68.3 (70.8) ; snout (o pelvic in.sortioii 3r).4 (3"). 5) and 
to pectoral insertion 31.4 (33.7). 

The following counts were made, respectively : Dorsal rays X-I, 22 
(X-I. 21) ; anal rays II, 9 (II, 9) ; pectoral rays ii, 16-ii, 17 (ii, 17-ii, 
17) ; pelvics always I, 5; vertical .scale rows counted above lateral lino 
•16 (46) and pores in lateral line 45 (46) ; scales from dorsal origin to 
lateral line 5 (5) and fi'oni base of first soft dor.^al ray to lateral lino 
5 (5) ; scales from lateral jiiie to anal (iriiiin 7 (7) : zig/ag .'-cale rows 
around caudal peduncle 17 (17). 

Snout blimtly roiuided, projecting a liltic in front of the mouth, tho 
latter inferior in [josition; lower jaw included: intciorbital space 
broad, a little convex, its width about equnl to Iciigdi of snout; an- 



130 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

terior profile from dorsal origin to between eyes nearly straight, or a 
very little convex; ventral profile curves to pelvic insertions, then 
nearly straight backward to anal origin; body compressed pos- 
teriorly; greatest depth of body at dorsal origin; eye 4:% in head, 
1% in interorbital space, and 2% in postorbital length of head; 
posterior nasal opening close to eye, larger than the anterior one; 
tip of lower jaw without barbels but with a median pit contain- 
ing two minute pores lying in this porelike depression, and laterally 
two pairs of pores as in panamensis; pores and lobes on front of snout 
as described for fanaineTisis ; anus a little over two-thirds closer to 
anal origin than to pelvic bases; tip of filament of pelvic soft ray 
reaching to anus ; pectoral fins reaching to opposite anus ; gill rakers 
moderately short, the longest equal to diameter of pupil ; preopercular 
spines numbering 9 or 10, none hooked downward, those dorsally 
smaller than those near lower angle of preopercle ; skull with the usual 
cavernous spaces as found in other genera ; least depth of caudal pe- 
duncle a little more than twice in its length ; teeth in villif orm bands in 
both jaws, the outer row of upper jaw a little enlarged; pseudo- 
branchiae well developed; scales ctenoid; lateral line broadly curved 
over pectorals, then running a straight course along midaxis of body 
posteriorly, and extending on the caudal fin; the fourth scale row 
below lateral line anteriorly is the first one continuing to base of caudal 
fin; second dorsal spine only slightly heavier (enlarged) than follow- 
ing spines ; second dorsal spine \y^ in second anal spine and reaching 
more than halfway to tips of third or fourth dorsal spines and 1% in 
postorbital length of head; second anal spine moderately enlarged, 
and not reaching to tips of soft rays ; pelvic spine equal to length of 
second dorsal spine ; distal margins of all fins a little rounded, that of 
caudal fin double truncate, with the middle rays longest. 

Color. — In alcohol, pale brownish above, lighter below; anal and 
spiny dorsal dusky; pelvic soft rays dusky distally, the filamentous 
ray white ; other fins very pale brownish ; peritoneum with numerous 
black pigment cells; the types are not well preserved and the colors 
have faded. 

ReTYiarhs. — This new species of OpMoscion may be separated from 
other Atlantic species of this genus by the foregoing key. 

Named hrasiliensis in reference to the country along whose shores 
the types were collected. 

OPHIOSCION MICROPS (Steindachner) 

Corvina microps Steindachner. Sitzb. Akad. Wiss. Wien [Ichth. Notizen No. 1], 
vol. 49, p. 6, pi. 2, fig. 2, 1864 (Guiaua). 

I have made measurements on two specimens from Para (M.C.Z. 
No. 4581), and the results are recorded below in hundredths of the 
standard length. Standard lengths in millimeters, 66.3 and 60.3. 



THREE NEW SCIAENID FISHES — SCHULTZ 131 

LenfiTth of head 32.1 and 33.'2; greatest depth of body 30.2 and 31.5; 
diameter of eye C.64 and G.47; length of snout 9.95 and 8.62; distance 
from front of upper lip to rear of maxillary 11.3 and 11,9; least pre- 
orbital width 4.98 and 4.48; postorbital length of head 18.5 and 19.1; 
bony interorbital space 10.4 and 11.3; length of caudal peduncle or 
distance from ba.se of last anal ray to raidcaudal fm base 26.8 and 24.9; 
least depth of caudal peduncle 9.95 and 10.1; length of base of second 
dorsal fin 33.2 and 35.6; length of anal fin base 12.5 and 12.6; length 
of longest or third dorsal spine 19.6 and 19.8; length of second dorsal 
spine 14.0 and 13.4 ; longest soft anal ray 10.6 and 21.5 ; length of second 
anal spine 18.1 and 18.6; longest pectoral ray 21.1 and 25.7; longest 
soft ray of pelvic fin 23.1 and 23.7 ; length of pelvic spine 12.4 and 13.3 ; 
tip of snout to dorsal origin 36.9 and 38.1 ; snout to anal origin 65.4 and 
66.8; snout to pectoral insertion 32.1 and 33.3; length of longest gill 
raker 2.71 and 2.65. 

The following counts were made : Dorsal rays XI-I, 19, and XI-I, 
21 ; anal rays II, 9 and II, 9 ; pectoral rays ii, 16-ii, 16 and ii, 17-ii, 16 ; 
pelvics always I, 5; gill rakers on first gill arch 11 + 1 + 19 and 
11 + 1 + 18; vertical scale rows above lateral line 51 and 50; scales 
from dorsal origin to lateral line 5 and 5, and from base of first soft 
dorsal ray to lateral line 4 and 4; scales from anal origin to lateral 
line 7 and 8; zigzag scale rows around caudal peduncle 18 and 18. 
Additional counts are recorded in table 1. 

This species has a very small eye, smaller than in any other species. 
Diameter of eye is contained in young li/> to 1% and in adults 2 to 
2% times in interorbital space. 

OPHIOSCION VENEZUELAE. new species 

FlOUKE 7 

Holotype. — U.S.N. M. No. 121749. one specimen, 139.5 nmi. in stand- 
ard length, collected by Leonard P. Schultz near mouth of Cano de 
Sagua, 25 km. north of Sinamaica, Venezuela, May 12, 1942. 

ParatypcH. — U.S.N.M. No. 121750, six specimens, 57 to 150 mm., 
collected with the holotype and bearing same data. 

Description. — Certain measurements were made, and these data, re- 
corded below, are expressed i)i liundrcdtlis of the standard length, first 
for the holotype and then for the thiee paratypes in i)aren(h('ses, re- 
spectively. Standard lengths in millimeters, 139.5 (68.8; 150; 139). 

I^^ngth of head 28.6 (30.5; 32.4; 30.2) ; greatest depth of body 30.1 
(27.0; 31.2; 30.9) ; diameter of eye 6.24 (7.12; 5.93; 6.11) ; length of 
snout 8.74 (8.14 ; 9.34 ; 8.63) ; distance from tip of snout to rear edge of 
maxillaries 13.1 (12.6; 13.0; 13.2) ; least width of preorbital 3.05 (3.63; 
4.13; 39.5) ; postorbital length of head IH.O (15.8; 17.9; 18.5) ; width of 
bony interorbital space 9.68 (9.16; 9.66; 10.3) ; length of caudal pe- 



132 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

duncle 25.1 (25.1 ; 24.3 ; 25.4) ; least depth of caudal peduncle 10.7 
(9.88; 10.9; 10.9) ; length of base of second dorsal fin 32.6 (32.5; 32.1; 
32.4) ; length of base of anal fin 11.0 (11.6; 11.7; 11.2) ; length of long- 
est dorsal spine 18.6 (21.1; 18.5; 19.3) ; length of longest soft dorsal 
ray — (13.1; —; — ; 13.2) ; longest soft anal ray 16.1 (17.0; — ; 14.7) ; 
length of second anal spine 16.3 (17.4; — ; 15.8) ; longest pectoral fin 
ray 25.2 (23.1; 22.7; 25.2); longest soft pelvic ray 13.0 (18.6; 12.3; 
13.9); length of pelvic spine 9.32 (11.5; 8.34; 8.85); longest or 
middle caudal fin rays 25.9 (26.9; 22.7; 25,5) ; distance from tip of 
snout to dorsal origin 37.1 (35.9; 38.5; 37.7); snout to anal origin 
67.2 (65.4; 65.1; 66.2); snout to pectoral insertion 32.6 (31.1; 32.2; 
31.6) ; snout to pelvic insertion 32.6 (30.5; 30.6; 80.9) ; length of long- 
est gill rakers on first gill arch 1.58 (2.76; 1.66; 3.22). 




Figure 7. — Opkioscion venezuelae, new species: Holotype (U.S.N.M. No. 121749). 

The following counts were made, respectively : Dorsal rays XI-I, 21 
(XI-I, 22; XII-I, 21; XI-I, 21; XI-I, 21; XI-I, 21; XI-I, 22); 
anal rays on all types II, 8; pectoral rays ii, 17-ii, 17 (ii, 16; ii, 
l7-ii, 17; ii, 17-ii, 17; ii, 16) ; pelvics always I, 5; number of vertical 
scale rows above lateral line 52 (53; 52; 54) ; scales from dorsal origin 
to lateral line 6 ( — ; 6; 6) and from base of first soft dorsal ray to 
lateral line 6 ( — ; 6; 6) ; scales from lateral line to anal origin 8 ( — ; 
8; 8) ; scales in a zigzag row around the caudal peduncle 19 ( — ; 
19; 19) ; number of gill rakers on first gill arch 9+1+16 ( — ; 10+1 
+ 18; 10 + 1 + 18; 9 + 1 + 16; 10 + 1 + 16). 

Head depressed forward but rounded dorsallj^, the interorbital space 
convex, broad, about equal to the snout; body compressed; anterior 
profile nearly straight but the dorsal contour curved, the ventral con- 
tour but slightly curved backward to anus; back highest at base of 
spiny dorsal fin ; eye about 2% in postorbital length of head, 14^ in 
interorbital space; posterior nasal opening rounded, slightlj^ larger 
than the anterior one; tip of lower jaw without barbels; anal origin 
equidistant between pelvic insertion and midcaudal fin base; pelvic 



THREE NEW SCIAENID FISHES — SCHULTZ 133 

fins reaching halfway to anus, tlie first soft ray ending; in a short fihi- 
ment; preuperole uith eight or nine short spines, the lowest one 
strongest but not hooked downward; caudal peduncle least depth 2V^ 
in its length; tips of pectoral fins reaching a trifle i)ast anus; teeth in 
jaws in bands, the outer row of upper jaw a little enlarged; pseudo- 
branchiae well developed; gill rakers short, not quite so long as 
pupil diameter; scales strongly ctenoid; lateral line curved over pec- 
toral fin, then running a straight course on caudal peduncle along 
its midaxis; fourth scale row below lateral line, anteriorly, the first 
one extending to base of caudal fin ; first dorsal spine rudimentary, 
second 2\-2 in third, the latter nearly as long as the fourth; second 
and eighth to eleventh and the next spine heavier than the third to 
seventh spines of dorsal fin ; fourth or longest dorsal spine about equal 
to postorbital length of head; distal margin of spiny dorsal fin trun- 
cate or a very little concave, that of soft dorsal probably a trifle rounded 
(the tips of the soft rays are lacking and this cannot be determined 
accurately) ; middle rays of caudal fin longest, edges of lobes more or 
less truncate to rounded (double truncate) ; distal margins of anal 
and pelvic fins a little rounded; pectoral fins somewhat pointed, the 
fourth branched ray from above longest. 

Color. — In alcohol the upper sides and back are grayish brown, 
white below; dorsal, anal, and pelvic fins dusky, more intensely pig- 
mented distally ; soft dorsal and cj^udal fins dusky; pectoral fin darker 
than other fins except tip of spiny dorsal; lower jaw and upper lip 
white; peritoneum white. In the smaller paratypes the dusky upper 
sides are broken up with several pale blotches, which appear to have 
a small cyst at their centers. 

Remar/cs. — This new species differs from all otlicr known species 
of OpJiioscion^ excei)t 0. adustus and 0. niicrops, both from the west- 
ern Atlantic, in having more numerous gill rakers, 1(> to 18 on lower 
part of first gill arch, but adustus has 28 soft dorsal rays and vene- 
zuelae only 22 to 24. O. microps has II, 9 anal rays and 0. venczuelae 
II, 8. The key will serve for distinguishing the seven species now 
recognized in the western Atlantic. 

Named venezuclae in reference to the country where the specimens 
were collected. 

OPHIOSCION I'UNCTATISSIMUS Mick and Hildcbrnnd 

OphioHcion punclntUiHimuH Mi;i;k and IIildkhuam), Mariiii' llshi'S of I'niiaina, vol. 

2. p. 044, pi. f«. WlTi (Cristobal, Torn Point, and Colon, I'anania). 
OphioMcion adunta Evkkmann and Maksii, Fislit's of Porto Kico, U. S. Fish Conim. 

Bull., vol. 110 (IDOO), pt. 1. p. 1:H>, lUOli ( Viftpics Island). 

I have examined the following specimens in the national collections: 
U.S.N.M. Nos. 817GG, the, holofype, and r>()l(;i and 120188, four 
specimens from Vieques Island off Puerto Kico. 



134 PROCEEDINGS OF THE NATIONAL MUSEUM vol.98 

U.S.N.M. No. 104297, one specimen, from Recife, Pernambuco, 
Brazil. 

U.S.N.M. Nos. 80765 and 80766, two paratypes, from Panama. 

OPHIOSCION PANAMENSIS, new species 
FiGXJEE 8 

Ophioscion adustus (in part) Meek and Hiujebrand, Marine fishes of Panama, 
vol. 2, p. 639, 1925 (Fox Bay, Colon, Panama). 

Holotype. — U.S.N.M. No. 122612, one specimen, 52 mm. in standard 
length, collected in Fox Bay, Colon, Panama, January 27, 1912, by 
Meek and Hildebrand. 

Paratypes. — U.S.N.M. No. 81204, three specimens, 31.5 to 43 mm. in 
standard length, from Fox Bay, Colon, Panama, January 5, 1911, 
Meek and Hildebrand ; U.S.N.M. No. 81205, four specimens, 23.3 to 
30.5 mm.. Fox Bay, Colon, Panama, March 31, 1911, Meek and Hilde- 
brand; U.S.N.M. No. 81207, one specimen, 33 mm., from Porto Bello, 
Panama, March 17, 1912, Meek and Hildebrand ; U.S.N.M. No. 81206, 
one specimen, 42 mm., collected along with the holotype and bearing 
same data; U.S.N.M. No. 128260, one specimen, 35.5 mm., from Fort 
Sherman, Canal Zone, Panama, collected March 3, 1937, by Dr. S. F. 
Hildebrand. 

Description. — Certain measurements were made, and these data, 
recorded below, are expressed in hundredths of the standard length, 
first for the holotype, then for a paratype in parentheses. Stand- 
ard lengths in millimeters 52 (42). 

Length of head, 33.3 (34.3) ; greatest depth of body 32.7 (32.2) ; 
diameter of eye 7.30 (7.86) ; length of snout 9.80 (9.76) ; distance from 
front of upper lip to rear tip of maxillary 10.4 (11.4) ; least preor- 
bital width 4.80 (4.76) ; postorbital length of head 17.9 (19.0) ; width 
of interorbital space 9.62 (10.2) ; length of caudal peduncle or dis- 
tance from base of last anal ray to midcaudal fin base 22.3 (23.1) ; 
least depth of caudal peduncle 11.2 (10.7) ; length of base of second 
dorsal fin 35.4 (35.7) and of base of anal fin 10.6 (11.9) ; longest dorsal 
spine 18.3 (16.4) ; longest soft ray of dorsal fin — (7.2) ; longest soft 
ray of anal fin 20.8 (18.6) ; length of second anal spine 18.8 (19.3) ; 
longest ray of pectoral fin 23.1 (22.4) ; longest soft ray of pelvic 21.7 
(25.0) and of pelvic spine 12.1 (12.6) ; longest midcaudal fin ray 30.0 
(31.0) ; length of longest gill raker 2.11 (2.38) ; distance from snout 
tip to dorsal origin 38.3 (39.3) and to anal origin 71.9 (67.4) ; snout 
to pelvic insertion 36.0 (35.6) and to pectoral insertion 34.6 (32.9). 

The following counts were made, respectively : Dorsal rays X-I, 20 
(X-1, 21) ; anal rays II, 7 (II, 7) ; pectoral fin rays ii, 17-ii, 17 (ii, 17-ii, 
17) ; pelvics always I, 5 ; scale rows above lateral line 51 (52) and pores 
in lateral line to midcaudal fin base 50 (50) ; scales from dorsal origin 



THREE NEW SCIAENID FISHES — SCHULTZ 135 

to lateral line 5 (5) and from base of first soft ray of dorsal to lateral 
line 5(5); t^cales from lateral line to anal origin 9 (8) ; zi<;za<j^ scale 
ro>Ys around caudal peduncle 19 (17). Additional counts are recorded 
in table 1. 

Snout bluntly rounded, projecting a little in front of mouth, the 
latter inferior in position, lower jaw included; interorbital space 
broaiK a little convex, its width about equal to length of snout ; anterior 
profile nearly straight from dorsal origin to between eyes or a trifle 
convex ; ventral profile a little convex anteriorly, then nearly straight 
to anal origin; body compressed posteriorly; greatest depth at dorsal 
origin; eye about 4V'> in the head, 21/5 m postorbital length of head, 
and 1V4 ill interorbital space; posterior nasal opening close to eye, a 



,„-^*!»"?^*'S!><> 



^ 



^ 



MiM. 



Figure 8. — Ophioscion panamensis, new species; Holotype (U.S.N.M. 122612). 

little larger than anterior one; tip of lower jaw without barbel, but 
with three pairs of pores, the median pair minute and in a porelike 
depression; tip of snout with two pairs of lobes, a pore lying between 
the middle pair of lobes and the outer pair being separated from the 
median lobes by a pore on each side, a third groovelike pore lying lat- 
erally to the outer lobe; then dorsally to the margin of the snout occur 
a median pore and another pore at each side; anus two-thirds closer to 
anal origin than to pelvic insertion; j)elvic fins not quite reaching to 
anus; the first soft ray of pelvics ending in a short filament; pectoral 
fins reafhing opposite tip of pelvics; gill rakers short, a little less than 
one-half the piij)il; preopercle with eight or nine small spines, those 
dorsally snmllei- th.m those near the lower angle of preopercle; none 
hooked downward; skull with the tisual ()|)en spaces or sinuses between 
the narrow bony biidges; least depth dl" caurjal peduncle l^^jo i" its 
iength; teeih in villiform bands in both jaws, the outer row of upper 
jaw slightly enlarged; i)seudol)ran<'hiae well developed; scales 
strongly ctenoid; lateral line broadly curved over pectorals, then run- 
ning a straight course along midaxis of body posteriorly, extending on 



136 PROCEEDINGS OF THE NATIONAL MUSEUM vol.98 

caudal fin ; the fifth scale row below lateral line anteriorly is the first 
one continuous to base of caudal fin ; second dorsal spine enlarged, its 
length ly^ in second anal spine, and reaching more than halfway to 
tip of third or fourth dorsal spine; second dorsal spine 1% in post- 
orbital length of head; second anal spine enlarged, not reaching to 
tips of soft anal rays ; pelvic spine a trifle shorter than second dorsal 
spine ; distal margins of dorsal, anal, and of paired fins a little rounded, 
that of the caudal fin double truncate, the midcaudal rays longest. 

Color. — In alcohol the body is brownish everywhere, paler brown 
ventrally; paired fins and anal and dorsal fins blackish, with the 
first spine in these fins, except pectorals, whitish ; tip of first soft ray 
in pelvic fins white, especially the filament ; soft dorsal and caudal fins 
brownish with numerous black pigment cells; undersides of head 
and breast pale ; lips pale ; peritoneum white. 

In the smaller specimens of this species the coloration of the median 
fins differs from the larger ones. At a standard length of 24 mm. 
the caudal fin is white, except for a few scattered brown pigment cells 
located near the center of the fin, the caudal fin base is abruptly dark 
brown with the pigment extending backward a little on middle rays ; 
the anal and dorsal fins have a brownish band extending across rays, 
with the margin of the fins white, and below this bar is another white 
area separating the brownish base of these fins from the brown band ; 
these fins gradually fill in with brown pigment so that at 42 mm. the 
fins are plain brownish. 

Bemarks. — This new species may be separated from other Atlantic 
species of Ophioscion by the key and traces down to O. adustus in 
Meek and Hildebrand's key to the species of Ophioscion in their 
"Marine Fishes of Panama" (vol. 2, p. 636, 1925) . 

Named panamcnsis in reference to the region where it has been 
collected. 

OPHIOSCION NASO (Jordan) 

Stelliferus naso Jordan, in Jordan and Eigenmann, Rep. U. S. Comm. Fish and 
Fisheries for 1S86, vol. 14, p. 395, 1889 (Cachiura, Brazil). 

I have made measurements on two of the types (M.C.Z. No. 4583) 
from Cachiura, and the results are recorded below in hundredths 
of the standard length. Standard lengths in millimeters, 75.5 and 
70.5. 

Length of head 31.1 and 30.5; greatest depth of body 31.8 and 28.4; 
diameter of eye 8.60 and 9.22 ; length of snout 9.27 and 9.22 ; tip of pre- 
maxillaries to rear of maxillary 10.1 and 9.78 ; least width of preorbital 
3.97 and 3.97; postorbital length of head 15.9 and 15.6; least width 
of bony interorbital 9.14 and 9.22 ; length of caudal peduncle (base of 
last anal ray to midbase of caudal fin) 25.0 and 24.4; least depth of 
caudal fin 10.6 and 10.8; length of base of second dorsal fin 31.8 and 



THREE NEW SCIAENID FISHES — SCHULTZ 137 

36.2; leno^th of anal fin base 12.4 and 12.1; lonf^th of longest dorsal 
or tiiird spine 22.3 and 21.0; longest soft dorsal ray 17.2 and — ; 
length of second dorsal spine 10.6 and 11.3; longest soft ray of anal 
fin 19.9 and 17.9; length of second anal spine 17.0 and 16.5; longest 
ray of pectoral fin 2r),2 and — ; longest soft ray of pelvic fins 21.2 
and 21.3; length of pelvic spine 12.6 and 12.3; longest or middle rays 
of caudal fin 28.2 and 28.4; tip of snout to dorsal origin 38.3 and 
37.2; snout to anal origin GG.2 and 68.8; snout to pectoral insertion 
31.8 and 31.9; longest gill raker 1.99 and 1.99. 

The following counts were made, respectively : Dorsal rays XI-I, 21 
and XI-I, 21; anal rays II, 8 and II, 8; pectoral rays ii, 16-ii, 16 and 
ii, 16-ii, 16; pelvics always I, 5; gill rakers on first gill arch 8 + 1 + 14 
and 8 + 1 + 14; vertical scale rows above lateral line 46 and 46; scales 
above lateral line at origin of spiny dorsal fin 4 and 4, and at base of 
first soft dorsal ray 5 and 5; scales below lateral line from anal 
origin to lateral line 8 and 8; zigzag scales around caudal peduncle 
18 and 18. 



u. t. «ov(iiNiiiNi rniNTiNs orricci i*4f 



PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 




SMITHSONIAN INSTITUTION 

U. S. NATIONAL MUSEUM 



Vol. 96 Wwhington: 1945 No. 3193 

THE ICHNEUMON-FLIES OF THE GENUS CRYPTANURA 
liRULLE, MAINLY TROPICAL AMERICAN 



P>V R. A. CUSIIMAN 



Because of BruUe's juistukcn idcu tluit the specimens on which he 
based the genus C ryptanxLva were fenuiles "with concealed ovipositors, 
this genus of ichneumon-flies was misunderstood until Roman (1910) 
identified it as the male of Polydeinis Cresson and synonymized the 
latter genus with Crijptanura. 

I'he genus dates from 1845, when the figure of Cryptanura nigripes 
Brulle was published in the atlas of Lepeletiers 'Tlistoire Natnrelle 
des Insectes," although tlie description did not appear until the fol- 
hjwing year. It is therefore a monobasic genus with nigripes Brulle 
as g<'notype. On this basis, and assuming that nigr/pcfi and .sfriafa 
Brulle are not congeneric, Viereck took exception to Roman's synon}'- 
mizing of Polyaenttfi Cresson with Cryptanura. Viereck's statement 
that these two species are not congeneric must have been based on the 
fact that Brulle does not state definitely that nigripes has the two 
small frontal horns characteristic of striata; but by inference he 
certainly does ascribe this character to nigripes^ for in the description 
of all the species that follow sfn'tifa he mentions only the cliuiinters 
by which they differ from it and from one another. 

Cresson and Cameron were in error in their inter])retati()ns of the 
genus, the species assigned to it by those authoi's Ijeing properly re- 
ferred to such genera as Trapr.zonaHs Szcpligeti, GUxlianus Cameron, 
I'liotocryptuft Viereck, and prol):ibly other genera. Roman and Brues 
iippear to be the only ones who have interpreted the genus correctly. 

624514—45 1 130 



140 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 9« 

Cryptanura is a large genus apparently confined in its distribution 
to the Western Hemisphere and there very largely to the tropical 
region, only two species being known to occur north of the Mexican 
border. 

Three species from the Old World Tropics have been referred to 
Polyaenus: cingvlatus Tosquinet from New Guinea, spiniferus Cam- 
eron from Borneo, and striatum Szepligeti from Formosa, but none 
appears to be properly referable to the genus. This is certainly true 
of cingulatu^ and spiniferus^ for I have been able to identify the 
latter definitely and the former without much question in the Baker 
collection. These si)ecies represent an apparently hitherto unde- 
scribed genus. It is described in an addendum to this paper. 

I have been unable to identify striatus Szepligeti, but its smooth 
and polished mesoscutum, longitudinally impressed propodeum with 
the apophyses apparently represented only by carinae, and elongate 
postpetiole would seem to exclude it from Cryptanura. Aside from 
the new genus mentioned above, to which striatus obviously does not 
belong, only one other Oriental genus is known to me that has two 
frontal horns. This is Ceratocryptus Cameron, in which striatus 
appears to be equally out of place. 

SPECIES WRONGLY REFERRED TO CRYPTANURA 

The following species described in Cryptanura by Cresson and 
Cameron do not belong to the genus. Unfortunately, most of them 
are unknown to me. The apparently proper status of the few that I 
have been able to place is indicated. 

{Cryptanura acolhua Cresson) =Glodianns acolhua (Cresson), new combi- 
nation. 

C albispina Cameron. 

C. cinctipcs Cameron. 

C. curtispina Cameron. 

C. delecta Cresson. 

C. fascia tipennis Cameron. 

C. incauta Cameron. 

C. interrupta Cameron. 

C. laticarinata Cameron. 

C. ornatipennis Cameron. 

(C pachymene Cresson) =Pliotocryptus pachymene (Cresson). 

C. pedicata Cameron. 

(0. sumichrasti Cvesson)—TrapezonaUs suniichrasti (Cresson), new combi- 
nation. 

Genus CRYPTANURA Brulle 

Cryptanura Brulle, Histoire naturelle des insectes : Hymenopteua, Atlas, pi. 41, 
fig. 6, 1845; vol. 4, p. 242, 1846.— Roman, Ent. Tidskr., 1910, p. 154.— Brues, 
Ann. Ent. Soc. Amer., vol. 5, p. 200, 1912. — Townes, Mem. Amer. Ent. Soc. 
No. 11, pt. 1, p. 288, 1944. 



ICHNEUMON-FLIES OF GENUS CRYPTANUIU — CUSHMAN 141 

Pulyacnus Crksson, Proc. Acad. Nat. Scl. riiihulolphia, 1873, p. 149.— Cameron, 
Kiologia Cvatrali-Aiufricana, Ilyuit'iioptfra, vol. 1, p. 244, 188G; Journ. Hoy. 
Agr. Coium. Soc. British Guiana, sor. 3, vol. 1, p. 16G. 1911.— St u.\nEm:KNKcnT, 
Genera insoctoruui, fasc. 7r>, p. G7, 19<,)8. — Sz^i-luieti, Ann. Mns. Nat. Hun- 
garici, vol. 14, p. 264, 1916.— Cushman, Proc. U. S. Nat. Mus., vol. 74, art. IG, 
p. 38, 1929. 

Polyacnidia Viekeck, Proc. U. S. Nat. Mus., vol. 46, p. 381, 1913. 

Hcsostcnus authors, part. 

Head from above transverse, the temples receding; frons with a 
carina medially and with two small horns, frequently arising from a 
common base; malar space distinct; clypeus strongly convex; eyes 
large and strongly convex; antenna in female frequently more or 
less distinctly thickened between middle and apex, the thickened por- 
tion flattened below. Thorax stout; epomia distinct and usually ex- 
tending to upper margin of pronotum, where they form carinate or 
conical projections; nolaulices deep and complete; scutellum small^ 
usually convex, rarely subconical or flattened ; propodeum with basal 
carina complete, apical carina usually represented only by two prom- 
inent apophyses, sometimes, especially in mule, distinct and witli 
apophyses less developed ; propodeum usually strongly rugose or 
transver.Ncly striate, rarely without sculpture, spiracles elongate; 
wings large; areolet small, complete, quadrangular, broadening to- 
ward apex, recurrent interstitial or somewhat antefurcal ; nervulus 
antefurcal; second discoidal cell broad at base; nervellus broken near 
bottom and perpendicular or weakly reclivous; logs long and usually 
rather slender; front tibia in female rarely slightly inflated. Abdo- 
men in female fusiform, in male small and narrow; spiracle of first 
segment far beyond middle; ovipositor sheath from a half to fully as 
long as abdomen; ovipositor subsagittate or swordlike at apex. 

Head and thorax ornamented with white or 3'ellow on a black or 
red ground; abdcjmen black and yellow or largely red; wings immacu- 
late hyaline or dilutely infumate. 

The color pattern of the head and thorax nearly throughout the 
genus is so similar as to constitute almost a generic character. In 
order to avoid much repetition a description of what may be called 
the normal or basic color pattern is gi\en here. In the specific descrip- 
tions only variations from this pattern are indicated. 

Ground color of head and thorax black, rarely red or partly red, 
with pale-yellow or whitish markings as follows: Oibilal ring, broad 
on cheek, much narrower or interrupted on upper temple and in malar 
space; face, clypeus (apical margin always dark), labrum, mandible 
basally, palpi, aninilus on antenna; pr()i)leura more or less; anterior 
and humeral margins of pronotum; meso.scutum either immaculate, 
or with a single spot on disk, or with paired lines along inner margins 
of lateral lobes, or with a nairow line on each lateral margin oppo- 



142 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 9e 

site tegulae, or with cuneiform markings on anterior margins of lat- 
eral lobes, or with combination of two or more of these; scutellum, 
its subtending carinae, and posterior margin of its f renum ; postscutel- 
lum and margin of its f renum ; tegulae ; subalar tubercles ; an oblique 
band on mesopleuron from near anterior margin to middle coxa; a 
larger or smaller spot on each side of mesosternum; upper division 
of metapleuron and its lower division largely, the latter with the 
ground color showing only below; and two broad marks posteriorly 
on propodeum embracing the apophyses, very rarely confluent ante- 
riorly. In the descriptions this color pattern is referred to as black 
(or red) with yellow or white markings and the exceptions noted. 

The abdomen is either largely red or the ground color is black, typi- 
cally with broad apical and lateral margins of tergites 1-7, lateral 
margins of tergite 8, and the petiole more or less yellow or whitish ; 
venter white with sternites darker. In the following descriptions this 
color pattern is referred to merely as black, the tergites margined 
with yellow and the exceptions noted. 

SPECIFIC CHARACTERS 

For the most part Cryptanura is a very homogeneous group with 
few striking structural characters that will serve to divide it into spe- 
cific groups, and it has been found necessary to depend to a consider- 
able extent on color in the construction of the following key to species. 

The form of the scutellum, the structure of the propodeum and 
pronotum, and the form of the ovipositor have been found useful as 
group characters, though the last, of necessity unisexual, does not 
appear to be accompanied by a companion character in the opposite 
sex. 

A considerable number of the described species have not been avail- 
able for study and have been omitted from the following key, but to 
assist in the identification of these species I have constructed a key to 
all the species, based largely on color. This will be found following 
the descriptions of the species examined. 

KEY TO SPECIES EXAMINED 

1. Scutellum strongly subconically elevated; metapleuron also with a tuberculate 

elevation 1. tuberculata, new species 

Scutellum and metapleuron not strongly elevated 2 

2. Apical carina of propodeum distinct medially at level of apophyses ; abdomen 

polished and unsculptured 3 

Apical carina usually absent, sometimes distinct in male, but then arching 
high above apophyses ; abdomen usually distinctly, finely sculptured— 5 

3. Abdomen and propodeum entirely red 2. dicostata, new species 

Abdomen and propodeum black and yellow 4 

4. Mesoscutum with yellow marginal markings, but without discal markings- 5 
Mesoscutum with paired discal markings, but without marginal markings. 

5. bicarinata, new species 



ICHNEUMON-FLIES OF GENUS CRYPTANURA — CUSHMAN 143 

5. Mesoscutal markings in furm of narrow marginal linos opposito togulae; 

torgite 2 yellow at apex, the j-ellow band broader medially; tegnlae yellow 

basally 3. quadrimaculata, new species 

Mesoscutal markings in form of cuneiform spots laterad of origins of nolau- 
lices; tergite 2 with a uniformly broad subapieal yellow band; tegulae 
entirely black 4. mediostrigosa, new species 

6. Ilunjeral margin of pronotum conicnlly prominent anteriorly; scutellum 

broad and transversely flat, with coarse, deep punctures; abdomen pol- 
ished, without trace of sculpture; front tibia in female subinflated; hind 
femur stout, not or barely two-thirds as long as tibia; pale markings of 

propodeum extending broadly forward to basal carina 7 

Disagreeing with all or nearly all above characters 10 

7. Abdomen black and yellow (or white) 8 

Abdnmen largely or entirely re<l 9 

8. Legs red, coxae marked with yellow; thorax reddish piceous and yellow. 

6. piceothorax, new species 
Legs and thorax black and yellow 7. planiscutellata, new species 

9. All coxae and front and middle femora black and yellow. 

8. politigaster, new species 
Hind coxae and aJl femora red 9. conica, new species 

10. EiKimia not extending upward to humeral margin of pronotum, the humeral 

njargin not at all carinate or tuberculate anteriorly ; ventrolateral carina 
of petiole distinct to base; second tergite with a median white spot at 

base 11 

EiK>mia forming a carinate elevation on humeral margin of pronotum ; ventro- 
lateral carina not distinct to base; second tergite without a median white 
spot 12 

11. Ht-ad and thorax red and yellow; antenna without white annulus; abdomen 

polished 10. ruficeps, new species 

Head and thorax black and yellow; antenna with white annulus; abdomen 
beyoml first tergite opaque shagreened, second and third tergites also 

punctate 11. septentrionalis, new species 

IZ M'soscutum with two yellow marks discally 13 

Mesoscutum with a single median spot, rarely flanked by small traces of 

yellow on inner margins of lateral lobes 16 

L3. Petiole and hind femur entirely black 12. apophysis, new species 

Pftiole and hind femur partly yellow 14 

H. I'ropodeal yellow spots not abruptly narrowed before apophyses, the latter 

long 15 

Proixtdeal yellow spots abiniptly narrowed before apophyses, the latter 

short 15. bilineata, new si>ecie3 

IT). Petiole yellow above, black bilow ; Iiiiul femur entirely black above. 

13. mexicana (Cresson) 
Petiole Itlack above, yellow below; hind femur with a narrow median yellow 

line above 14. orizabensis (Canjeron) 

10. Thorax red and yellow 16. rufa, new species 

Tlt'irax black and yellow 17 

17. Proiwideul spots each with a narrow forward extension, sometimes (male) 

confluent medially 18 

ProiKxleal .sjwt.s either not extending forward or not abruptly narrowed 
before ap^iphyses 22 

18. Abdomen black and yellow 19 

Abdomen largely or entirely red 20 



144 PROCEEDINGS OF THE NATIONAL MUSEUM vol. m 

19. Front and middle femora red ; hind femur red ( $ ) or entirely black (S) ; 

hind coxa red ( 9 ) or black ( $ ) with a yellow spot above. 

17. spinaria (Brull6) 

All femora black and white, hind femur white below, black above, with a 

narrow median white line; hind coxa white below and above, black 

on each side 18. lineatifemur, new species 

20. Hind coxa black and white 19. coxata, new species 

Hind coxa largely or entirely red 21 

21. Humeral margins of pronotum prominent, subtuberculate anteriorly; post- 

petiole red . 20. boliviensis, new species 

Humeral margins of pronotum merely carinate anteriorly, not prominent; 
postpetiole piceous, margined with yellow 21. isthmus, new species 

22. Ovipositor nearly as long as abdomen, stout, much deeper toward apex than 

at base, not sagittate at apex 23 

Ovipositor slender, its upper and lower margins parallel except at the sagit- 
tate apex, usually much shorter than abdomen 25 

23. Abdomen red 22. excalibur, new species 

Abdomen black and white 24 

24. Hind coxa largely and femur red 23. acinaces, new species 

Hind coxa yellow and black ; femur black or piceous above, yellowish below. 

24. propinqua (Cresson) 

25. Front and middle femora entirely red 25. pretiosa (Viereck) 

Front and middle femora piceous posteriorly 26 

26. Hind leg very slender, femur apparently nearly 8 times as long as deep; 

temples in dorsal view very strongly receding and slightly concave. 

26. gracilipes, new species 

Hind leg stouter, femur apparently not more than 6 times as long as deep; 

temples flat or weakly convex or concave 27 

27. Temple at middle of eye distinctly more than half as broad as short diame- 

ter of eye and very weakly convex 28 

Temple not or barely half as broad as short diameter of eye and fiat or weakly 
concave 29 

28. Head in side view with occipital carina very nearly parallel to posterior 

margin of eye ; apical margin of clypeus weakly curved. 

27. gracilis, new species 

Occipital carina and posterior margin of eye distinctly divergent below; 

apical margin of clypeus perfectly straight 28. genalis, new species 

29. Hind femur rather slender, at least 6 times as long as deep and more than 

three-fourths as long as tibia ; flagellum in female much thickened beyond 

middle and flattened below, the joints there strongly transverse 30 

Hind femur rather stout, distinctly less than 6 times as long as deep and 
less than three-fourths as long as tibia ; flagellum in female only slightly 
thickened and flattened beyond middle, the joints there weakly trans- 
verse 32 

30. Occipital carina nearly parallel to posterior margin of eye ; ovipositor sheath 

hardly as long as abdomen beyond first tergite 29. variegata (Brull6) 

Occipital carina and posterior margin of eye distinctly divergent below; 
ovipositor sheath nearly as long as abdomen 31 

31. Temple at middle of eye distinctly less than half as broad as short diameter 

of eye ; flagellum very slender at base, first joint more than 6 times as long 

as thick at middle 30. paranensis, new species 

Temple about half as broad as short diameter of eye ; flagellum stouter, flrst 
joint not 6 times as long as thick at middle. 

31. tenuiterebrata, new species 



ICHNEUMON-ILIES OF GENUS CHYPTANURA — CUSHMAN 145 

32. Face entirely yt'Uow ; frttnt and middle coxae piceous behind, yell*)w in 

front, metathorax not at all red 32. incerta (Cresson) 

Face with longitudinal iniprt'ssions broadly black ; front coxa largely 
piceous, middle coxa entirely red; nielai)lenron below and nietasternum 
red 33. maculifrons, new siiecies 

1. CRYPTANURA TUBERCULATA, new speciei 

Evidently clctsely allied to .scutellaris (Szopligeti) and possibly syn- 
onymous with tliat species. However, it seems unlikely that 8zepli- 
geti would have failed to mention tlie very evident tuberculiform 
metapleura charaeteristic of the present species if his species were of 
similar structure. 

Female. — Length 19 nun., antenna 17 mm., ovipositor sheath 8 mm. 

Head in dorsal view with temples very slightly concave, occipital 
carina flangelike, sinuate at lower extremity; frontal horns very 
small; scrobes very deep; a distinct, radiately rugose impression sur- 
rounding the ocelli except behind; postocellar line hardly as long as 
ocellocular line, stemmaticum longitudinally rugose; eyes parallel 
within, faintly sinuate opposite frons; face with a median subhemi- 
spherical elevation flanked on each side by a transversely rugose im- 
pression; clypeus smooth, with scattered punctures, roundly convex 
with a broad reflexed margin, broadly truncate; malar space very 
nearly as long as basal width of mandible ; antenna only slightly thick- 
ened beyond middle, 38-jointed. Thorax polished, with notaulices, 
margins of mesoscutum, sternaulices, and groove along posterior mar- 
gin of mesopleuron foveolate; scrobe of pronotum, upper anterior por- 
tion of mesopleuron, metapleuron, and sides of propodeum obliquely 
striate; mesopleuron below and sternum sparsely punctate; humeral 
margin of pronotum smooth, anteriorly tuberculate; scutellum very 
strongly elevated, subconical, in profile with its posterior slope deeply 
concave ; metapleuron with a high rounded tubercle in the middle ; legs 
very long and slender, hind femur reaching distinctly beyond apex 
of abdomen and fully eight times as long as deep; coxae polished and 
sparsely punctate. Abdomen almost exactly as long as head and 
thoi-ax, rather narrow, very finely shagi'eened, subopacjue beyond first 
tergite; i)etiole distinctly depressed; ovipositor ratiier stout, becoming 
gradually slightly deeper toward apex, where it is swordlike rather 
than subsagittate. 

Head and thorax black with yellow maculation (see description of 
color patern, p. 141), abdomen ferruginous, legs ferruginotis and 
])arlly yellow and piceous; wings stibhyaline, with blackish venation; 
orbital ring interrupted on upper temples and in mahir sjiace; facial 
impressions and outline of clypeus black; anmilus on antenna embrac- 
ing flagellar joints (5)r)-12( b'J) ; two yelU)w lines on disk of mesos- 
cutum; apex and anterior face of the elevated part of scutellum black; 
sternum yellow only along sternaulices; lower division of metapleuron 



146 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

with only the tubercle and its apex yellow ; stripes on propodeum ex- 
tending only a short distance basad of apophyses. Front coxa behind 
and all femora behind piceous, this color reduced on hind femur to a 
narrow stripe on the inner side and the apex; front legs otherwise yel- 
lowish, as are also the middle coxa, tibia, and tarsus, the hind tibia 
except at apex, and a stripe on inner side, which are blackish, and the 
hind tarsus. Ovipositor sheath black, 

Tyfe locality. — St. Laurent du Maroni, French Guiana. 

Types. — A holotype and a paratype, both females, taken at the above 
locality by Audoit in 1862. The type is in the Paris Museum ^ and the 
paratype in the U. S. National Museum (No. 57080) . 

The paratype is a little larger than the type but is otherwise like it. 

2. CRYPTANURA DICOSTATA, new species 

Apparently related to shnilis (Szepligeti) and perhaps the same 
but having the apical carina of the propodeum more or less distinctly 
developed, a character not mentioned by Szepligeti. 

Female. — Length 12 mm., antennae 11 mm., ovipositor sheath 4 mm. 

Temples very narrow and strongly receding, flat; occipital carina 
neither especially prominent nor sinuate at lower extremity; vertex 
and frons deeply concave, ocelli distinctly below level of superior 
tangent of eyes, frontal horns small; face nearly flat, transversely 
rugulose and very minutely shagreened, subopaque; clypeus strongly 
convex, with very narrow reflexed margin, polished, with sparse 
punctures; malar space about two-thirds as long as basal width of 
mandible, finely opaque; head elsewhere polished; postocellar line 
and diameter of an ocellus about equal and much shorter than ocel- 
locular line; antenna 33-jointed, slender, only slightly thickened and 
flattened beyond middle. Thorax anteriorly subopaquely sculptured, 
posteriorly polished and almost without sculpture ; humeral margin of 
pronotum smooth, anteriorly angulate, scrobe striate; mesoscutum 
punctate, prescutum transversely striate on each side next to the 
notaulices, lobes flattened; scutellum convex, polished, with a few 
punctures ; mesopleuron striatopunctate above, punctate below, smooth 
posteriorly; sternum punctate; metapleuron polished and sparsely 
punctate, with a few rugae posteriorly; propodeum smooth and 
polished, more or less roughened medially between carinae; apical 
carina distinct and straight between apophyses; basal carina only 
slightly curved medially ; basal area not at all defined ; legs long, hind 
femur about six times as long as deep and reaching slightly beyond 
apex of abdomen; coxae polished. Abdomen distinctly longer than 
head and thorax, polished, unsculptured, rather slender ; first segment 
without trace of dorsal carinae, petiole slightly broader than deep, 

1 All specimens indicated as being in, or received from, the Paris Museum are retained 
In the U. S. National Museum for the duration of the war. 



ICHXEUMOX-FLIES OF GENUS CRYPTANURA — CUSHMAX 147 

I>ostjietiolo barely as ^vi(le as lonp:; second teririto as ]oujz ^^ fii'^t and 
about twice as lon<j: as broad at base; ivopositor sheath shorter than 
abdomen beyond first segment; ovipositor slender, of even depth 
throughout, apex weakly subsagittate. 

Head and anterior portion of thorax black with yellow markings 
(see description of color pattern, p. 141) ; thorax posteriorly, mesoster- 
num, abdomen, and legs ferruginous; antennal annulus embracing 
flagellar joints (4) 5-11 (12), two narrow yellow streaks on disk of 
mesoscutum, scutellum yellow only in basal angles and at apex, meso- 
pleuron yellow except the prepectus, the impression below tubercle, 
and an oblique streak running down from this, which are black; 
upper division of metapleuron yellow tinged with ferruginous; propo- 
deum, lower section of metapleuron, and sternum, except prepectus, 
ferruginous; front coxae anteriorly and all tibiae and tarsi yellow; 
front and middle femora piceous below and behind; wings yellowish 
hyaline, venation dark brown; abdomen entirely ferruginous, sheath 
biack. 

7^l/pe locality. — Kamakusa, British Guiana. 

Type!i. — Four females, the holotype and three paratj'pes in the 
U. S. Natitmal Museum (No. 57058). 

The type and two paratypes were taken at the type locality by H. 
Lang. The third paratype is from Port-of-Spain, Trinidad. 

One specimen of what is probably the male of this species was taken 
by W. M. Mann at Cavinas, Beni, Bolivia, during the Mulford Bio- 
lr)gical Exploration of 1921-22. It has the mesopleuron and sternum 
almost entirely yellowish stramineous, the antennal annulus embrac- 
ing fl.igellar joints (5) 6-19 (20), and the apical carina of propodeum 
only faintly indicated medially. It is excluded from the type series. 

8. CRYPTANURA QUADRIMACULATA. new species 

Apparently closely' allied to cctypa (Cresson), from the description 
of which it differs in its possession of yellow lateral lines on the meso- 
scutum. Also Ciesson makes no mention of the presence, in his 
species, of the apical carina of the propodeum. 

Frmalr. — Length 13 mm., antennae (broken), ovipositor sheath 4.5 
nun. 

Temples flat, short and very sharply receding, occipital carina 
moderately prominent, bent abruptly inward at its lower extremity 
to meet the hypostomal carina far back from base of mandible; vertex 
and frons concave, lateral ocelli slightly below superior tangent of eyes, 
frontal horns very acute and arising from common base; postocellar 
line and diameter of an ocellus equal and distinctly shorter than ocel- 
locuiar line; eyes large and bidging, weakly fonvergent lielr»\v ; 'i\n-\.' :md 
rlyjieus minutely shagreened, sub()j)aque, face laterally and dyiu'Us 

C24.'>14— 45 2 



148 PROCEEDINGS OF THE NATIONAL MUSEUM vol.98 

sparsely punctate, face dorsally irregularly striate; clypeus strongly 
convex, in profile subnasute; malar space hardly two-thirds as long 
as basal width of mandible. Thorax rather slender, generally polished ; 
pronotum striate in lower part of scrobe, its upper anterior margin 
only weakly tumid, smooth, carinately angled; mesoscutum sparsely 
punctate, rugose in posterior middle, prescutum more densely punctate 
and transversely striate anteriorly along notaulices, lobes flattened; 
mesopleuron largely and sternum sparsely punctate, space below tu- 
bercle more or less striate ; scutellum convex, with scattered punctures ; 
metapleuron very sparsely punctate and with a few vertical striae 
posteriorly; propodeum with sparse, coarse punctures basally and 
more or less distinctly longitudinally striate medially behind basal 
carina, basal area obsoletely defined, basal carina bent sharply for- 
ward medially, apical carina distinct between apophyses ; legs slender 
but hind femur less than two-thirds as long as tibia and not reaching 
apex of abdomen ; coxae elongate, polished, and sparsely punctate, are- 
olet narrow. Abdomen longer than head and thorax, polished, with 
scattered punctures on basal tergites ; first tergite narrow, petiole not 
depressed, dorsal carinae absent, postpetiole barely as broad as long, 
with a median impression between spiracles; second tergite as long 
as first and nearly twice as long as broad at base ; sheath nearly as long 
as abdomen beyond first tergite ; ovipositor slender, of nearly uniform 
depth, and subsagittate at apex. 

Black and yellow (see description of color pattern, p. 141) ; cheeks, 
frontal orbits, an annulus beginning on flagellar joint 6 (missing 
beyond joint 8) yellow; anterior margin of pronotum black except 
for a small spot ; humeral margin entirely black ; narrow lines on lateral 
margins of mesoscutum above tegulae, only basal angles and apex of 
scutellum, tegulae basally, and mesopleuron except prepectus largely 
yellow, the last confluent with mark on sternum, metapleuron except 
suture between upper and lower divisions and a dash near apex en- 
tirely yellow, large yellow spots in basal lateral areas of propodeum, 
apical markings of propodeum with triangular projection anteriorly, 
apical bands on basal tergites triangularly broadened medially; 
petiole yellow above, black below ; venter yellow. Legs yellow, with 
front and middle coxae behind, hind coxa within and medially above, 
middle trochanter and femur behind, hind trochanter largely, and 
femur except a yellow streak on each side black or piceous; wings 
hyaline, venation blackish. 

Male. — Essentially like female, the frontal horns much longer. 

Type locality. — San Bernardino, Paraguay. 

Types. — A female holotype, an allotype, and a female paratype in 
the U. S. National Museum (No. 57059). All were collected at the 
above locality by K. Fiebrig. 

The paratype female is essentially like the type. 



ICHNEUMON-FLIES OF GENUS CRYPTANURA CUSHMAN 149 

4. CRYPTANURA MEDIOSTRIGOSA, new species 

Female. — Length 14 mm., antenna 12 mm., ovipositor sheath 4.5 mm. 

Differs from the above description of qiuulrimaculata principally 
as follows: Mesopleuron polished, with a small area of rather coarse 
j)unctiires anteriorly; metapleuron rather coarsely* punctate-rugose 
over most of its surf race; propodeum polished basally, longitudi- 
nally striate medially between the transverse carinae and in i)etiolar 
area; hind femur fully two-thirds as long as tibia and fully reaching 
apex of abdomen; abdomen polished and entirely without sculpture; 
petiole depressed; postpetiole fully as broad as long, flat between 
spiracles. 

Color pattern as in quadrim-amdata except frontal orbit broader 
and extending over top of eye; antennal annulus beginning on joint 
5; anterior margin of pronotum entirely black, humeral margin with 
a white mark at the angle; mesoscutum with cuneiform markings 
extending bacJvward from origins of notaulices; tegulae entirely 
black; mesopleuron and sternum and metapleuron and sternum stra- 
mineous rather than yellow, this color including also most of the pre- 
pectup; iJropodeum dorsally yellow except for a cruciform black 
mark with its crossbar on the basal carina ; yellow bands of abdomen 
subapical and of nearly uniform width; front and middle legs yel- 
low, with femora posteriorly and apical half or more of tarsi black- 
ish; hind leg stramineous, trochanter and femur black above. 

Male. — Essentially like female. 

Type locality. — Barro Colorado Island, Canal Zone. 

Types. — Two females collected at the type locality by S. W. Frost, 
the holotype on March 10, the paratype on March 22, 1937; and one 
male allotype taken by A. Busck on March 12, 1912, at Alhajuela, 
Canal Zone. All are in the U. S. National Museum (Xo. 57UC0). 

S. CRYPTANURA BICARINATA, new fpecies 

Though related to qu/idrhnaculata in its possession of the distinct 
apical carina and in its highly polished abdomen, this species is very 
tlistiiict, especially in the minute frontal horns, stouter body, and 
ditferi'ut arrangement of color. It seems uidikely that this can be 
championi (Cameron), also from Panama, for it is apparently much 
stouter and has the black on (he inner side of the hind fcnuir rather 
than on the ui)per side. Moreover, Cameron does not mention the 
I)resenco in his species of the apical carina of the propodeum. 

Fnnah. — Li'ngth 13 mm., an(enna«' 12 nun., ovipositor sheath 5.5 
mm. 

Temples flat, narrow, sharply receding, occipital ciuiiia rather 
prominent, ils lower extremity curving slightly to join the hyposlo- 
mal carina back from the base of the mandible; vertex and frons con- 



150 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

cave, ocelli slightly below superior tangent of tlie eyes ; frontal horns 
obsolete, their common base distinct ; eyes large, bulging, very faintly 
convergent below ; face rather flat, with a transversely striate impres- 
sion on each side of middle, the upper striae extending across middle, 
laterally opaque»shagreened, medially polished ; clypeus polished and 
sparsely punctate, in profile strongly convex ; malar space hardly two- 
thirds basal width of mandible; antennae 32-jointed, only very 
slightly thickened, and flattened beyond middle. Thorax rather stout, 
polished and sparsely punctate except as noted below ; humeral mar- 
gin of pronotum smooth, carinately angled anteriorly, scrobe striate 
below ; lobes of mesoscutum rather flat, prescutum transversely striate 
along notaulices; scutellum weakly convex; mesopleuron anteriorly 
and metapleuron in lower posterior angle obliquely striate; propo- 
deum with basal area defined, basal carina arching forward medially, 
apical carina distinct between the very stout, compressed apophyses, 
middle of propodeum both before and behind apical carina obsoletely 
transversely rugose, and with a few coarse rugae radiating from the 
apical margin ; legs rather stout, hind femur hardly five times as long 
as thick, less than two-thirds as long as tibia and barely reaching - 
apex of abdomen; areolet rather broad, its lower side distinctly 
angled by the strongly antefurcal recurrent. Abdomen polished, 
stout, petiole distinctly depressed, postpetiole broader than long, dor- 
sal carinae absent ; second tergite shorter than first and barely a half 
longer than broad at base ; ovipositor sheath about as long as abdomen 
beyond first tergite, ovipositor rather slender, of uniform depth, and 
weakly subsagittate at apex. 

Black and yellow (see description of color pattern, p. 141) ; orbital 
rings of unusually uniform width behind eyes; annulus embracing 
flagellar joints (5) 6-10 (12) ; yellow of anterior margin of pronotum 
reduced to two small spots near middle ; two small streaks on disk of 
mesoscutum ; scutellum with an elongate black mark in basal middle ; 
mesopleuron entirely yellow; yellow spots in basal lateral areas of 
propodeum and the apical bands extending forward nearly to basal 
carina; petiole yellow both above and below; broad subapical bands 
and lateral margins of tergites 2 and 3 yellow, the color extending 
inward at base of second ; remaining tergites entirely and venter yel- 
low. Front and middle legs yellow, with femora and middle tro- 
chanters piceous posteriorly ; middle coxa piceous on each side above ; 
hind coxa similarly marked but more reddish beneath; trochanter 
stained with piceous; femur reddish on outer side, piceous within; 
tibia and tarsus yellow. Wings hyaline, venation black. 

Type locality. — Trinidad Kio, Panama. 

Type. — One specimen taken by August Busck, March 16, 1912. It 
is in the U. S. National Museum (No. 57061). 



ICHNEUMON-FLIES OF GENUS CRYPTANURA — CUSHMAN 151 
6. CRYPTANURA PICEOTHORAX, new species 

Female. — Longtli 1-2 iniu.. rtntcnniie 8.5 inin., ovipositor shealli 
3.5 mm. 

Temples narrow, flat, and sharply receding; occipital carina prom- 
inent, especially below, where it joins the hypostomal carina not far 
from the base of the mandible; vertex nearly straight across except 
for a groove at each side of ocelli; ocelli at level of superior tangent 
of eyes; frons moderately concave, slightly tumid laterally, horns 
prominent on a common base; eyes moderately large and somewhat 
bulging, weakly convergent below; face with a rather deep striate 
impression on each side of middle, the middle roundly convex and 
with sparse, coarse punctures; clypeus sculptured like middle of face, 
strongly convex, in profile subnasute; malar space flat, nearly as long 
as basal width of mandible; antennae stout, slightly thickened and 
flattened near apex, 29-jointed. Thorax stout, generally coarsely 
punctate to striatorugose; pronotum with humeral margin smooth^ 
conically produced anteriorly, scrobes coarsely striate; mesoscutum 
coar.-^ely punctate, prescutum somewhat rugose, lobes rather flat; 
scutellum flat and coarsely punctate above, apex rather abrupt and 
impuMctate; frena of both scutellum and postscutellum coarsely 
foveolate; pleura and propodeum coarsely striatorugose; sternum 
coarsely punctate; basal areas partly smooth; basal median area 
l)artly defined, basal carina nearly straight; legs rather stout, hind 
femur little more than two-thirds as long as tibia and not extending 
to apex of abdomen; hind coxa coarsely punctate above; areolet short. 
Abdomen broad, polished; petiole strongly depressed, post petiole 
much broader than long, with a weak elevation on each side of middle 
between spiracles, but without dorsal carinae; second tergite a little 
shoiter than first and only about a third longer than broad at base; 
sheath sliorter than abdomen beyond first tergite; ovipositor rather 
thick, in profile rather slender, a little deeper near apex, the apex not 
at all sagittate. 

Head and thorax black to rcdtlisli piceous, both with yellow mark- 
ings (see description of color pattern, p. MI); amiuhis emliiacing 
flagellar joints (5) 6-9 (10) ; single spot on disk of mesoscul urn ; only 
apex (tf scutellum yellow; most of nies(»pleuron and sternum and 
nietaj)leiiron except suture sepai'al ing its two divisions yellow ; pro[)o- 
deMJ marking extending broadly forward to basiil caiina, but con- 
stricted before apophyses. Legs ferruginous, coxae and trochanters 
partly yellow. Wings hyaline, veins blacU, stigma brown. Abdomen 
lilack with white markings; petiole, except vent rally, white ; three spots 
t lan.-vei-sely airange(] at base of second teigite and a bioad subapical 
band also white; the extreme a[)ex i)lack. 



152 PROCEEDINGS OF THE NATIONAL MUSEUM vol. se 

Type locality. — Baragua, Camagiiey, Cuba. 

Types. — Two females, the holotype taken at light June 4, 1932 
(Christenson), the paratype at Sole, Cuba, March 2, 1925 (George 
Salt). Both are in the U. S. National Museum (No. 57063). 

The paratype is smaller, with the sculpture less heavy and the dark 
color of the thorax more reddish. 

7. CRYPTANURA PLANrSCUTELLATA, new species 

Very similar in form, structure, and color pattern to piceothorax 
but at once distinguishable by the color of its thorax, which is not at 
all red, and of the legs, which are black and yellow. Differs further 
from piceothorax as follows : 

Female. — ^Length 12 mm,, antennae 8.5 mm., ovipositor sheath 
5.5 mm. 

Sculpture of thorax, especially metapleuron and propodeum, in- 
cluding the basal areas, more reticulate-rugose than striatorugose ; 
scutellum less abruptly sloping at apex ; postpetiole and tergites 2 and 
Z coarsely, sparsely punctate, especially laterally. 

Antennal annulus longer, extending to flagellar joint 13; pale mark- 
ings of abdomen yellow rather than white; legs to apices of femora 
black and yellow, tibiae and tarsi pale stramineous ; front and middle 
coxae yellow, black behind, hind coxa yellow with base below and 
streaks on upper and inner sides black; trochanter yellow, basal joint 
above and apical joint at apex more or less black, hind trochanter with 
basal joint largely black; front and middle femora yellow, black be- 
hind; hind femur yellow on outer and inner sides, black above and 
below. 

Type locality. — Puerto Castilla, Honduras. 

Type.— One female dated March 26, 1924. It is in the U. S. National 
Museum (No. 57062). 

This may be the same as compacta (Cresson). 

8. CRYPTANURA POLITIGASTER, new species 

This species is similar in structure and in color pattern of head 
and thorax to the preceding two species but differs from both in its 
red abdomen and in other details. 

Female. — Length 11 mm., antennae 9 mm., ovipositor sheath 3 mm. 

Temples rather long, slightly but distinctly concave, sharply reced- 
ing, occipital carina high, especially at lower extremity, where it 
passes over a prominence in the posterior margin of the cheek, and 
joins the hypostomal carina rather close to the base of the mandible; 
cheek transversely striate along carina, vertex flat with a shallow 
groove next to ocelli; frons deeply concave, tumid next to eyes, ob- 
liquely striate above, horns small but elevated on a high common base ; 



ICHNEUMON-FLIES OF GENUS CRYPTANURA — CUSHMAN 153 

eyes large, somewhat buliriiig, weakly convergent below; face nearly 
smooth laterally and medially with a rugose impression on each side 
of middle, the rugae extending across middle at top; clypeus smooth 
with large scattered punctures, nasute in profile; malar space flat and 
fully as long as basal witlth of mandible; antenna rather stout, weakly 
thickened and flattened near apex, 30-jointed. Thorax stout; humeral 
margin of pronotum smooth, conically prominent anteriorly, scrobe 
striate; mesoscutum coarsely punctate, lobes rather flat, present um 
very low anteriorly; scutellum sloping from base to apex, polished, 
with a few large punctures; frena coarsely foveolate; mesopleuron 
longitudinally striate below tubercle, obliquely and indistinctly so 
below, elsewhere polished and sparsely punctate, prepectus rugu- 
lose-punctate, sternum polislied and sparsely punctate; metapleunm 
coarsely rugose above, polislied and punctate below, as is also its upper 
division; propodeum largely rugose, more or less transversely so 
behind apophyses, basal median area defined, basal carina curved 
medially; legs rather slender but not long, hind femur barely two- 
thirds as long as tibia and not reaching apex of abdomen; areolet 
short. Abdomen broad, polished, with a few large punctures at sides 
of postpetiole and second tergite; first segment stout, petiole flattened 
above but not distinctly depressed, postpetiole much broader than 
long, without trace of dorsal carinae; second tergite nearly as long 
as first, but barely a third longer than broad at base ; ovipositor sheath 
distinctly shorter than abdomen beyond first segment; ovipositor of 
uniform depth; apex weakly subsagittate. 

Head and thorax black with yellow markings (see description of 
color pattern, p. 141) ; annulus embracing flagellar joints (5) 6-11 
(13); anterior margin of pi-onotum medially and humeral margins 
before tegulae black; spot in center of mesoscutum; scutellum yellow 
only at apex; mesopleural band divided into a large spot anteriorly and 
a smaller one posteriorly; a large spot on each side of sternum; both 
npj)er and lower divisions of metapleuron largely yellow; propodeal 
markings extending broadly forward to carina but constricted before 
apophyses. Front and middle legs yellow, with their coxae, tro- 
chanters, and femora black behind; hind coxa black and yellow, tro- 
chanter and femur ferruginous, tibia and tarsus yellow. Wings 
hyaline, venation black. Abdomen fei'iiiginous; petiole and apex of 
fii-st tergite yellow, postjM'fioh! ferruginous and black; stcrnites pale 
ferruginous margined wilh whitish. 

Type locality. — Kurrenabaque, Itif) Beni, I'olivia. 

Type. — One female, taken in October by W. M. Mami on the 
Mul ford P.iological E.\i)loration of 191^-22. It is in the U. S. National 
Museum (No. 570C5). 



154 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 9» 

9. CRYPTANURA CONICA, new species 

Similar to politigaster but distinguishable at once by its much more 
strongly flexed occipital carina and by its entirely red femora and 
hind coxae. 

Female. — Length 12 mm., antennae (tips broken off), ovipositor 
sheath 3.5 mm. 

Temples rather long, slightly concave, and sharply receding; occi- 
pital carina high, especially below, where it passes over a strong angu- 
lation in the posterior margin of the cheek; vertex concave, ocelli dis- 
tinctly below superior tangent of eyes; frons moderately concave, 
with rugae radiating from ocelli, horns small but elevated on a broad 
common base ; eyes bulging, very weakly convergent below ; face with 
a shallow impression on each side of middle, medially coarsely rugose, 
laterally smooth; clypeus with scattered coarse punctures, in profile 
subnasute; malar space fully as long as basal width of mandible, flat; 
antennae distinctly thickened and flattened beyond middle. Thorax 
stout, largely coarsely sculptured; pronotum with humeral margin 
transversely striate, conically produced anteriorly, scrobe coarsely 
striate; mesoscutum coarsely but not densely punctate, lobes, espe- 
cially prescutum anteriorly, flattened ; mesopleuron striate, the sculp- 
ture changing to punctation toward lower posterior angle, sternum 
polished and sparsely punctate; scutellum roundly sloping from base 
to apex, polished, with sparse punctures, frena foveolate; metapleu- 
ron obliquely punctatostriate, its upper division smooth and sparsely 
punctate; propodeum polished before carina, coarsely rugose behind, 
the rugosity becoming transverse on apical slope, basal median area 
defined, basal carina curved forward medially; legs rather slender 
but not very long, hind femur barely two-thirds as long as tibia and 
not reaching apex of abdomen, hind coxa polished, with sparse punc- 
tures; areolet narrow. Abdomen rather narrow, polished; petiole 
flat above but not depressed, postpetiole much broader than long, with 
no trace of dorsal carinae; second tergite about as long as first and 
about a half longer than wide at base ; sheath much shorter than abdo- 
men beyond first tergite; ovipositor slender, of uniform depth, and 
subsagittate at apex. 

Head and thorax black with yellow markings (see description of 
color pattern, p. 141) ; orbital ring broadly interrupted on temples; 
annulus on flagellar joints (5)6-10(13) ; anterior margins of prono- 
tum entirely and humeral margin posteriorly black; mesoscutum im- 
maculate ; apex of tegulae piceous ; mesosternum immaculate ; yellow 
marking of lower division of metapleuron reduced to a streak in the 
dorsoposterior half; propodeal markings extending from apex to 
basal carina and narrowing and divergent from apophyses to carina. 
Legs ferruginous, tibiae and tarsi and front and middle coxae ante- 



ICHNEUMON-FLIES OF GENUS CRYPTANURA — CUSHMAN 155 

riorly yellow. AVings hyaline, venation black. Abdomen ferrugi- 
nous, petiole and apex of postpetiole stramineous. 

Type localitj/. — Kamakusa, British Guiana. 

Type. — One female taken by H. Lang. It is in the U. S. National 
Museum (No. 57066). 

10. CKYPTANUKA RUFICEPS, new spcciea 

This and the next following species differ from all others known to 
me in lacking any trace of angulation of the tumid humeral margin 
of the pronotum; and the present species is unique in the genus for 
its red head and is very unusual in its lack of a pale annulus on the 
antenna. 

Female. — Length 13 mm., antennae 10.5' mm., ovipositor sheath 
4.5 mm. 

Temples short, distinctly convex, sharply sloping, occipital carina 
rather high, especially at lower end, hypostoraal carina very high; 
vertex straight across, ocelli elevated above general level; frons not 
deeply excavated, horns very small, on a broad, low, common base; 
eyes not bulging; parallel within; face subpolished, with coarse punc- 
tures, especially in sliallow impressions on each side of middle, median 
portion in profile nearly as strongly convex below as is clypeus, 
which is not so strongly convex as usual and is sculptured like the 
face: malar space a little shorter than basal width of mandible; cheeks 
in front view slightly convex, in side view fully twice as broad as 
malar space; antenna slightly thickened and flattened beyond mid- 
dle. Thorax robust, sculpture mostly coarse rugosity; pronotum with 
scrobe coarsely striate, epomia strong, but terminating below the 
smooth humeral margin, which is less strongly tumid anteriorl}' than 
posteriorly; mesoscutum with lobes very weakly convex and notaulices 
very shallow, polished and sparsely, coarsely punctate; scutellum 
strongly convex but not prominently so, polished, with a few punc- 
tures, frena foveolate; mesopleuron largely coarsely rugosostriate; 
sternum coarsely punctate; metapleuron and propodeum, except its 
punctate basal areas, transversely striate, basal median area not de- 
fined, basal carina curved forward me^lially; apophyses unusually 
short and obtu.se; legs rather slender, hind femur three-fourths as 
long as tibia and reaching apex of abdomen, hind coxa polished and 
punctate; areolet broad. Abdomen stout, i)olislied; petiole distinctly 
flepressed, ventrolateral carina strong, with a foveolate groove above 
it, postpetiole much broader than long, with low rounded elevations 
repre.sentmg the dorsal carinae; second tergite about a fourth longer 
than broad at base; ovipositor sheath not quite so long as abdomen 
beyond first tergite, ovipositor rather stout, subsagittate at apex. 

824514— 4!5 3 



156 PROCEEDINGS OF THE NATIONAL MUSEUM vol, 66 

Head and thorax rufous with yellow markings (see description of 
color pattern, p. 141) ; orbital ring not abruptly narrowed on temples; 
facial impressions, clypeal suture, and margin of clypeus ferruginous ; 
scape rufous, flagellum entirely black; mesoscutum with a single 
median spot; scutellum with a rufous spot in basal middle; meso- 
stemum and both upper and lower divisions of metapleuron largely 
rufous; propodeal markings each with a narrow tongue-shaped ex- 
tension before the apophyses and reaching nearly to basal carina. 
Antenna black, scape rufous, without white annulus; small areas in 
rufous coloring of head and thorax more or less piceous; legs entirely 
ferruginous ; wings yellowish hyaline, venation black. Abdomen black 
and white ; three white spots at base of second tergite, the middle one 
large and broadly transverse, the lateral ones small and longitudinal ; 
venter mostly white ; sheath black. 

Type locality. — Pico Turquino, Cuba. 

Types. — A holotype and four paratypes, all females, collected by 
S. C. Bruner and C. H. Ballou, July 10-20, 1922, at elevations ranging 
from 2,900 to 6,000 feet. They are in the U. S. National Museum (No. 
57067). 

11. CRYPTANURA SEPTENTRIONALIS, new species 

Related to unifjceps in the lack of the angulation of the humeral 
margins of the pronotum and in the color pattern of the abdomen, but 
distinct from that species in the black and white head and thorax 
and in many details of structure and color. 

Fem-ale. — ^Length 15 mm., antennae (broken), ovipositor sheath 
4,5 mm. 

Differs from ruflceps principally as follows : Frons obliquely striate 
above, horns not on a common base ; postvertex densely, temples, cheeks, 
and sides of frons more sparsely, punctate; face rugose except later- 
ally, less strongly convex medially; malar space much shorter than 
basal width of mandible; occipital and hypostomal carinae weaker; 
sculpture of thorax largely coarse punctation, densest and coarsest on 
mesoscutum and metapleuron, humeral margin of pronotum coarsely 
punctate ; propodeum posteriorly irregularly transversely rugose, scu- 
tellum polished ; notaulices deep ; apophyses longer ; abdomen beyond 
first tergite opaque shagreened, second and third tergites and post- 
petiole at sides punctate ; ventrolateral carinae of petiole subobsolete. 

Differs from ruflceps in color as follows: Head and thorax black 
and white ; face entirely white ; antennal annulus embracing flagellar 
joints (5) 6-11 (12) and underside of scape white: propodeal white 
marks narrowly confluent across median line anteriorly ; coxae white, 
the front and middle ones more or less black behind, the hind coxa 
black on outer and inner sides toward base and with a median black 
stripe above, trochanters largely white, legs otherwise ferruginous, 
with tarsi paler. 



ICHNEUMON-FLIES OF GENUS CRYPTANURA CUSHMAN 157 

Type locality. — Cleveland, Ohio. 

Types. — One female (the holotype) captured September 19, 1932, by 
Frank D. DeGant; one female paratype, St. Louis, Mo., June 29, 1938; 
and one female paratype received from H. A. Scullen, Corvallis, Oreg., 
but probably eastern. A\\ are m the U. S. National Museum (No. 
57068). 

This is the second species of this tropical genus to be found in the 
United States. 

12. CRYPTANURA APOPHYSIS, new specieii 

Distinct from all tlie other species known to me in its extraordinarily 
large propodeal apophyses. It is also almost unique in its entirely 
yellow front and middle legs and black hind femur. 

Female. — Length 15 mm., antennae (missing), ovipositor sheath 
5.5 mm. 

Temples flat to slightly concave, long, sharply receding; occipital 
carina very high, but becoming abruptly lower just before its junction 
with the very high hypostomal carina; vertex concave, top of ocelli 
below superior tangent of eyes; frons with short rugae radiating 
from ocelli, deeply concave below, horns short, stout, not on a com- 
mon base; eyes very large, bulging, very weakly convergent below; 
face medially elevated, with an impression on each side of middle, 
opaque .shagreened, with sparse punctures, obliquely rugose below 
antennae; clypeus subnasute, subpolished, with a few coarse punc- 
tures; malar space nearly as long as basal width of mandible, straight 
in front view. Thorax distinctly compressed ; pronotum with humeral 
margin transversely striate, carinately angulated anteriorly, scrobe 
striate; mesoscutum subpolished and rather densely and coarsely 
punctate, lobes low, notaulices deep and narrow; scutellum strongly 
I on vex, polished, very sparsely punctate; mesopleuron subpolished, 
-par.sely punctate below, scrobe polished below and striate above; 
mesosternum more densely punctate; upper division of metapleuron 
coarsely punctate, lower division coarsely obliquely rugosostriate ; 
propfMleum rugose laterally and between carina and apophyses, 
otherwise largely polished, apopl»y.ses very large, long, conical, nearly 
twice as long as their distance from basal carina; legs very slender, 
hind femur fully eight times as long as deep and reaching beyond apex 
of abdomen; areolet elongate. Abdomen s](!nder, linely shagreened 
and suliopaque beyond first tergite; petiole ahnost exactly .square in 
cioss-section. {)ost[)etiole nearly as long as broad; second tergite fully 
two-thirds longer than l)road at base; sheath as long as abdomen 
beyond first tergite; ovipositor rather stout, distinctly subsagittate at 
apex. 

Black and yellow (see description of color pattern, p. 141) ; orbital 
ring nearly interrupted in upper temple and in malar space; two 



158 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 96 

narrow lilies on disk of mesoscutum ; a large spot anteriorly on meso- 
pleuron and a smaller one before middle coxa ; yellow of lower division 
of metapleuron reduced to a large spot in the middle of its upper 
margin; propodeal spots extending only a short distance anterior to 
apophyses ; front and middle legs except their coxae at base behind, 
a large spot on upper side of hind coxa, hind tibia except narrowly at 
base and broadly at apex, and probably hind tarsi (though these are 
missing) yellow; petiole entirely black; wings yellowish hyaline, 
venation blackish. 

Male. — Essentially like female, but more highly polished ; the pro- 
podeum with striae radiating from bases of apophyses, which are 
somewhat more slender than in female; postpetiole fully as long as 
broad and second tergite fully twice as long as broad at their junction ; 
legs even more slender. 

The antennae, though not entire, exhibit a yellow annulus beginning 
on the eleventh flagellar joint, and the hind tarsus is entirely yellow. 

Type locality. — Colombia. 

Allotype locality. — Huascaran, Peru. 

Types. — One of each sex, the type female, having no further data ; 
the allotype taken September 21, 1911, by C. H. T. Townsend. Both 
are in the U. S. National Museum (No. 57069) . 

13. CRYPTANURA MEXICANA (Cresson), new combination 

Mesostenus mexicanus Cresson, Proc. Acad. Nat. Sci. Philadelphia, 1873, p. 157, 
female, male (part). 

Cresson's type series was not all of the same species, for a male 
eotype in the National Collection is referable to orizdbensis (Cam- 
eron). A female eotype agrees with Cresson's description, and ap- 
parently is the true meooicana. 

Three additional specimens are before me as follows : Argas, 
Panama, April 28, 1911 (A. Busck) ; Costa Rica and Santa Marta, 
Colombia, 1852 (Fontanier), the last belonging to the Paris Museum. 

Female. — ^Length 14 mm., antennae 14 mm., sheath 5.5 mm. 

Temples flat or slightly concave, occipital carina high, slightly 
sinuate below; vertex weakly concave, ocelli a little below superior 
tangent of eyes ; postocellar line much shorter than ocellocular line ; 
frons moderately concave, with a few rugae radiating fi'om the 
ocelli, horns short and broad, not on a common base ; eyes large, bulg- 
ing, weakly convergent below; face convexly elevated medially, pol- 
ished and sparsely punctate, with a transversely striate impres- 
sion on each side of middle; clypeus strongly convex but not subnasute, 
polished and sparsely punctate; malar space nearly as long as basal 
width of mandible, in front view straight; antenna about 38- jointed, 
strongly thickened and flattened below beyond middle. Thorax 
stout; humeral margin of pronotum transversely striate, carinately 



ICHNEUMON-FLIES OF GENUS CRYPTANURA — CUSHMAN 159 

aiighnl anteriorly, scrobe striate; niesoscutuiii polished, sparsely 
t^oarseiy puiu-late. notaulices deep and narrow; scutellum narrow, 
convex, polished, with a few punctnres; mesopleuron obliquely striate 
alx)ve, the striation chan<rin<T £rradually to punctation below, sternum 
punctate; upper division of metapleuron punctate, lower division 
coarsely, obliquely rugosostriate ; propodeuni with basal median area 
obsoletely defined, basal carina curving forward medially, apophyses 
long and slender, basal areas punctate and posteriorly more or less 
rugose, space between carina and apophyses with oblique striae con- 
verging toward middle, posterior face more coarsely, transversely 
striate; legs lt)ng and slender, hind femur reaching apex of abdomen, 
hind coxa sparsdly, coarsely punctate; areolet elongate. Abdomen 
beyond fii-st tert^ite finely shagreened, subopaque; petiole not de- 
pressed, post petiole broader than long, with median and lateral shal- 
low impressions; sheath a little longer than abdomen beyond first 
tergite ; second tergite about a half longer than broad at base; oviposi- 
tor subsagittate at apex. 

Black with yellow markings (see description of color pattern, 
p. 141) ; orbital ring interrupted in upper temples; annulus embracing 
flagellar joints (5) 0-10 (12) ; two elongate marks on disk of meso- 
scutum ; scutellum yellow only at apex, mesosternum largely yellow : 
lower division of metapleuron very largely yellow; jiropodeal mark- 
ings extending forward nearly to carina; petiole yellow above, black 
below. Legs yellow ; front and middle coxae and femora behind, base 
of hind coxa and a broad stripe above, and apical joint of hind tro- 
chanter and the fennir above black. Wings hyaline, narrowly, weakly 
infumate apically, venation blackish, 

I have not seen a male of this species, but according to Cre^ison il 
differs from the female in being more slender, with antennae and 
legs longer, the knees and the apex of the hind tibia black. 

U. CRYPTANLRA ORIZABENSIS (Cameron), new combination 

SfiHoxtenua mcTinimiH Ckkhson, Proc. Acad. Nat. Sfi. Pliiladelpbia, 1873, p. 1157, 

mule (part). 
I'olyafnun orizabrnxiH Camkkun, Hlologla Centrall-Aiuericaiia. Hyuienoptera, vol. 

1, 188G, p. 240, female. 

As stated above, the National Museum male colype of iiwxicana 
(Cre.s.son) belongs to this species. A female, also from Mexico, from 
the C. F. Baker collection is also before me. 

The female differs from that of tncx-i/iana in structure principally 
as follows: Occipital carina not sinuate below; postocellar line nearly 
as long as ocellocular line; frontal horns longer and more acute; 
sculpture at middle of proi)odeum irregularly transverse rugosity; 
ovipositor and sheath distinctly .shorter; areolet shorter, with recur- 
rent interstitial. 



160 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

Differs from mexicana in color as follows : Mesosternum yellow only 
along sternaulices ; propodeal markings extending only a little before 
apophyses; hind coxa black above with a large oval yellow spot; 
basal joint of hind trochanter largely black, apical joint yellow, hind 
femur black above but with a narrow median yellow line, hind tibia 
somewhat blackish or brownish at base and apex ; petiole black above, 
yellow below. 

The male is very much like the female but has the frontal horns 
longer and more acute and the hind tibia more broadly black at apex. 

15. CRYPTANURA BILINEATA, new species 

Very similar to the two preceding species, but distinct from both 
in the form of the propodeal markings, which extend forward to the 
carina but are abruptly narrowed before the apophyses; in the less 
distinctly thickened and flattened antennae; in having the frontal 
horns set rather high on a common base; in the broader and more 
weakly convex scutellum; and in the short, obtuse apophyses. It is 
like Tnexicana in general in the color of the legs but has the hind coxa 
colored as in orisabensis; the petiole is, as in mexicana, yellow above 
and black below; and the mesosternum, as in oriBobensis, is yellow 
only along the sternaulices. In the long postocellar line and in the 
sculpture of the propodeum it is more like ori^abensis and in the 
length of the ovipositor and in venation of the wings it agrees with 
mexicana. 

Type locality. — Cuernavaca, Morelos, Mexico. 

Type.—On& female in the U. S. National Museum (No. 57064), 
taken by E. G. Smyth. 

16. CRYPTANURA RUFA, new species 

Distinct from all other species known to me and apparently from 
all other described species in the arrangement of color, black and 
yellow head, red and yellow thorax (not black dorsally), and entirely 
red abdomen. 

Female. — ^Length 14 mm., antennae 12 nmi., ovipositor sheath 
5 mm. 

Temples weakly convex, occipital carina not high, with a shallow 
notch just before its junction with the hypostomal carina ; vertex flat, 
ocelli as high as superior tangent of eyes ; f rons rather weakly concave, 
with rugae radiating from ocelli, horns short, stout, rather widely 
separated ; eyes weakly convergent below ; face polished and sparsely 
punctate, weakly convex medially, with a shallow, transversely striate 
impression on each side of middle; clypeus polished and sparsely 
punctate, subnasube in profile; malar space three-fourths as long as 
basal width of mandible ; cheeks in front view weakly convex. Thorax 
robust ; humeral margin of pronotum transversely striate, carinately 
angled anteriorly, scrobe sparsely striate ; mesoscutum rather densely 



ICHNEUMON-FLIES OF GENUS CRYPTANURA — CUSHMAN 161 

punctate, notaulices deep and narrow, lobes rather flat; scutellum 
convex, polished, and obsoletcly punctate; mesopleuron obliquely stri- 
ate, this sculpture changing below and posteriorly to rather sparse 
punctation. sternum more densely pun'-tate; upper division of meta- 
pleuron polished and sparsely punctate, lower division coarsely, 
obliquely rugosostriate ; propodeum polished and punctate before and 
transversely rugosostritite behind carina, posterior fare transversely 
striate, apophyses moderately long and slender; legs rather stout, hind 
femur hardly reaching apex of abdomen, hind coxae polished and 
sparsely punctate. Abdomen ■^ubpolished, inconspicuously shagreened 
beyond first tergite; petiole slightly depressed, postpetiolc much 
broader than long, with faint rounded elevations between the spiracles ; 
second tergite about a third longer than broad at base; sheath shorter 
than abdomen beyond first tergite, ovipositor rather stout, subsagit- 
tate at apex. 

Head black (see description of color pattern, p. 141) with orbital 
lings interrupted in upper temple and in malar space; facial impres- 
sions and outline of clypeus piceous. Thorax ferruginous, slightly 
stained with piceous in postscutellar frena and in speculum and with 
yellow markings; single spot on disk of mesoscutum; scutellum yellow 
only at apex: upper divisioii of metapleuron and dorsal half of its 
h'wer division yellow; propodeal markings including the apophyses 
and two small spots just in front of and mesad of these. Front and 
middle legs yellow, with front coxa, trochanter, and femur piceous 
behind, middle coxa ferruginous with yellow spot in front; hind leg 
ferruginous witli tibia and tarsus yellow. Wings hyaline, venation 
blacki.sh. Abdomen entirely red. 

Type locality. — Estero de Sao Paulo, Brazil. 

Typea. — A female holotype and two female paratypes tnken in 1910 
by E. R. Wagner. Tlie holotype and one paratjqoe are in the Paris 
Museum, and the second paratype is in the U. S. National Museum 
(Xo. 57081). There is also a female with l)roke,n antennae in the Paris 
Mu.seum, taken in 1010 at Cliaeo de Santiago del Estero, Argentina. 

17. f RYPTANURA SPINARIA (Brullc) 

Mr.nnHtenuft Mpinariuif BRt:T,l.f;, Hislnjrf njitun-llr' dos ins<'<-fos, Hyiin'iioplern, vol. 
4. p. 227, 184f5, feraalo. 

^fC80fltfmuH alhnpirtus rBERBON hiul Smith), I'ror. Ent. Sor. rhiladelphin. vnl. 3. 
p. 312. 1804. male. 

}friiO!iU-nu.<< drinii nrrnsia Dali.a Toiire. CntnloRus liyiiiciiojifornninT, i>. HIO. lfK)1~ 
02. 

I'Dlfinriix spiti'irifix SriiMlKOKK M' iii. (IfiK-rn iiist'rtnruiii. f)iH<'. 7"t. p. C,S. IJtilS. 

\frsoKtrnuH spituiriiiH ViESKCK, in Sinitli, Inserts nf Now Jorsoy. p rC^O, 1f>1(t. 

^fc808tenid^a (Vnlynrnun) spinaria Vikkdi;k, The HyiiHMioptcra <>r wnsp-liko In- 
sects of Connef.tlcut, p. 320, 1917. 

Pnlyncnun Kpinnriim fxTSiiMAN, Joiirn. ^^■nsllinKtn^ Acnrl. Sri., vol. 15. p. 301. 10"Jr» : 
Pror. U. S. Nat. Mus., vol. 74. nrt. 10. p. .3S. fl«s. U. 3fc, W, 1020. 

Crupl">iiirn ^pinnrin Tdwnks, M<>m Aiikt. Knf. Soc , No. 11, jit. I. j). L'MS. 1011. 



162 PROCEEDINGS OF THE NATIONAL MUSEUM vol,. M 

I have described this North American species too recently and too 
fully to make further discussion necessary here. It may well be men- 
tioned, however, that the mesoscutum frequently has, in addition to 
the single median yellow spot, traces of tlie yellow lines along the 
inner margins of the lateral lobes and sometimes those along lateral 
margins next to the tegulae. 

18. CRYPTANURA LINEATIFEMUR. new species 

Fennale. — Length 14 mm., antennae 11 mm., sheath 5 mn\. 

Except that it has a single median spot on the mesoscutum instead 
of two yellow lines and that the propodeal markings extend narrowly 
forward to the basal carina, this species is very similar in color and in 
large part in structure also to the three species mexicana^ orizabensis, 
and hilineata. The hind legs are colored exactly like those of oriza- 
iensis, as is the mesosternum, while the abdomen is colored like that 
of mexicana. The temples are not at all concave, the cheeks are more 
convex, tlie vertex is flat, the occipital carina is lower, the antennae 
are shorter, the petiole is thicker, with the lateral carinae, both dorsal 
and ventral, more distinct, and the ovipositor longer ; otherwise very 
similar to oiizabensis. 

Type locality. — La Caja, near San Jose, Costa E-ica. 

Types. — Two females, the holotype and .i paratype, taken at the 
above locality by M. Valeria on July 1, 1931, at an altitude of 900 
meters, and one female paratype captured by Schaus and Barnes at 
Cayuga, Guatemala. All are in the U. S. National Museum (No. 
55071). 

The Costa Rican paratype is smaller and the Guatemalan paratype 
larger than the type, but otherwise they are similar. 

Mesostenus veraepacis Cameron, which seems to be a Ci^ptarmra, 
appears to be very similar to this species but differs in that the hind 
femur is entirely black above without the yellow median line. 

19. CRYPTANURA COXATA, new species 

Female. — Length 16 mm., antennae 12 mm., ovipositor sheath 
5.5 nam. 

Temples flat, short, and very strongly receding, occipital carina 
moderately high, somewhat higher and slightly sinuate below ; vertex 
nearly straight across ; f rons rather deeply concave above, with striae 
radiating from ocelli, horns short and broad, close together but not on 
a common base ; eyes slightly bulging, weakly convergent below ; face 
polished and sparsely punctate, somewhat rugose below antennae, 
impressed on each side of middle; clypeus strongly convex, polished, 
sparsely punctate ; malar space about two-thirds basal width of mandi- 
ble; cheeks in front view weakly convex; antenna 32-jointed, dis- 
tinctly broadened and flattened beyond middle. Thorax stout; hu- 
meral margin of pronotum transversely striate, carinate anteriorly. 



ICHNEUMON-FLIES OF GENUS CRYPTANURA — CUSHMAN 163 

scrobe striate; mesoscutuni polished aiul sparsely punctate, lobes some- 
what flattened; scutellum convex, polished, very sparsely punctate; 
upper division of nietapleuron polished and sparsely punctate, lower 
division coarsely, obliquely ru^osostriate; pro[)odeuni transversely 
striate behind apophyses, before which it is irregularly rugose, basad 
of carina polished medially and rugosopunctate laterally, apophyses 
rather long and slender; legs rather stout, hind femur hardly reach- 
ing apex of abdomen. Abdomen stout, subopaque shagreened beyond 
lii-st tergite; petiole depressed, postpetiole much broader than long 
wiih two rather prominent elevations between spiracles; second ter- 
gite hardly a third longer than broad at base; sheath much shorter 
than abdomen beyond first tergite, ovipositor rather stout, subsagittate 
at i\[K'x. 

Head and thorax black with yellow markings (see description of 
color i)attern, p. 141) ; orbital ring interrupted in upper temple; annu- 
his em!)racing ilagellar joints (5)6-10(12) ; a single spot in center of 
]iifsoscutum; large spot on each side of sternum; propodeal markings 
abrui)tly narrowed but not constricted before apophyses, and curving 
tr)ward each other just behind the carina. Front and middle legs 
yellow, the coxae, trochanters, and femora black behind; hind coxa 
yellow below and in middle above, black above on outer and inner sides, 
trochanter black basally, reddish apically, femur ferruginous, tibia 
and tarsus yellow ; wings yellownsh hyaline, venation blackish. Abdo- 
men ferruginous, first segment black with petiole dorsally and apical 
margin yellow. 

J/,/A. — Much smaller than female, with apical carina complete and 
.strongly arched forward medially, the apophyses reduced to highei 
elevations in the carina, rugosity of thorax, except in potiolar area, 
largely replaced by punclation; abdomen much smaller and narrower, 
legs relatively longer and more slender; clypeus entirely yellow 
except apical margin; scape beneath; annulus in flagellar joints 
(10)11-17(18); propodeal markings confluent anteriorly; all abdo- 
minal segments beyond first more or less blaikish basally. 

7'j/pe local if y. — Tiinidad Kfo, Panama. 

TyjH'.s. — 'I'wo females and one male, the liolot ype an<l all<)lyi)e, taken 
March 27 and 20, 1912, by August Busck, ami a i^arntype female by 
\V. M. Mann at Ixianias, Bolivia, in Deeembei- 1!>21, on the Mulford 
liiological Ivxploration. All are in the U. S. National Museinn (No. 
.07070). 

20. CRYPTANURA BOLIVIENSIS. new upcritu 

Very closely related to coxdfa, frcjm which it can at once be distin- 
guished by the entirely red hind coxa and trochanter and first teigile. 

Fnnalc. — Length ir» mm., antenna 11.;") mm., ovipositor sheath T) mm. 

Differs further fi'om roxdia as follow^: 'I'eniples longei- and less 
-harply receding; humeral margins of pronotum subtuberculate 



164 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

anteriorly; lateral lobes of mesoscutum nearly impunctate; scutellum 
impunctate; thorax laterally much less strongly striate, mesopleuron 
mostly punctate rather than striate; abdomen narrower; petiole not 
depressed, postpetiole not especially broad, though somewhat broader 
than long, second tergite more than a half longer than broad at base. 

Antennal annulus embracing flagellar joints (4) 5-11 (14) ; pro- 
podeal markings strongly constricted before apophyses and not curving 
mesad anteriorly ; middle coxa and trochanter ferruginous behind. 

Type locality. — Rosario Lake, Rogagua, Bolivia. 

Type. — One female taken in November 1921 by M. R. Lopez, on the 
Mulford Biological Exploration. It is in the U. S. National Museum 
(No. 57072). 

21. CRYPTANURA ISTHMUS, new species 

Female. — ^Length 15 mm., antenna 12 mm., ovipositor sheath 5 mm. 

Structurally almost identical with coxata., but diflFering in color as 
follows : Mesosternum yellow only along sternaulices ; propodeal mark- 
ings constricted basad of apophyses and not curving toward each other 
anteriorly; hind coxa ferruginous, with an indistinct yellow spot at 
base above; petiole stramineous, postpetiole piceous with apical mar- 
gin yellow, second tergite obsoletely yellow just before apex. 

Type locality. — Trinidad Rio, Panama. 

Types. — Two females, the holotype and a paratype, taken by August 
Busck at the type locality on March 16 to 19, 1912; and one female 
paratype taken by C. T. Greene at Ajicon, Canal Zone, May 17, 1926. 
They are in the U. S. National Museum (No. 57073). 

22. CRYPTANURA EXCALIBUR, new species 

This and the next following two species and probably Mesostenus 
platyurus Brulle form a group distinct from the rest of the genus in 
the long, stout ovipositor which in side view is almost clavate, becoming 
gradually much deeper toward the apex. Brulle's species is not known 
to me, but it may be distinguished from the present species by its red 
propodeum. 

Female. — Length 16 mm., antennae 13 mm., ovipositor sheath 9 mm. 

Temples very weakly convex, rather short, occipital carina high, 
especially just before joining hypostomal carina; vertex straight 
across; frons rather shallowly concave, with rugae radiating from 
ocelli, horns short, thick, separate; eyes very weakly convergent 
below; face subpolished, sparsely punctatorugose above, somewhat 
elevated medially, with a shallow impression on each side of eleva- 
tion; clypeus strongly convex, polished and sparsely punctate; malar 
space two-thirds basal width of mandible ; cheeks in front view con- 
vex; antenna 36-jointed, very slightly thicker and flattened toward 
apex. Thorax stout; humeral margin of pronotum transversely stri- 
ate, carinately angled anteriorly, scrobe striate; mesoscutum subpol- 



ICHXEUMON-FLIES OF GENUS CRYPTANURA — CUSHMAN 165 

ished and coarsely punctate, lobes weakly convex; scutellum stron<j;Iy 
convex, polished, nearly inipunctate; niesopleuron obliijuely striate 
above, rather densely punctate below, sternum similarly punctate; 
upper division of metapleuron polished and sparsely punctate, lower 
division coarsely, obliquely rugosostriate; propodeum striate behind, 
the striations becoming somewhat confused above, basad of carina lat- 
erally rugose, medially polished and sparsely punctate; apophyses 
rather stout; legs stout, fenmr hardly reaching apex of abdomen. 
Abdomen very finely shagreened and subopaque beyond first tergite; 
petiole slightly depressed, postpetiole distinctly broader than long, 
a shallow impression on each side above spiracle; second tergite more 
than a half longer than broad at base; ovipositor sheath nearly as long 
as abdomen, ovipositor very stout, nearly twice as deep near apex as at 
base. 

Head and thorax black with yellow markings (see description of 
color pattern, p. 141); occipital ring nearly or quite interrupted on 
temple; annulus on flagellar joints (5)6-11(13) ; scape entirely black; 
mesoscutum with a single median spot ; mesosternum yellow only along 
sternaulices; propodeal markings extending broadly forward from 
apophyses but not reaching carina. Front and middle legs yellow, 
the coxae, trochanters, and femora black behind ; hind leg ferruginous, 
a small spot at base of coxa above and the tibia and tarsus yellow. 
Abdomen ferruginous, postpetiole apically and laterally yellow; 
sheath black. 

Type locality. — Trinidad Rio, Panama. 

Types. — Three females, all collected by August Busck, the holotype 
on March 29, one paratype on March 23, 1912, at the type locality, and 
a paratyjje on June 3, 1007, at Tabernilla, Canal Zone. They are in 
the U. S. National Museum (No. 57074). 

The paratypes are slightly smaller than the type but otherwise 
much the same. 

13. CRYPTANURA ACINACES, new ipecies 

Female. — Length lo mm., antennae (broken), ovipositor sheath 
8 mm. 

Structure very similar to that of excalibiir, l)ut abdomen blaclc with 
yellow markings. From the above descript ion of fxra/J/fur the pi'esent 
species differs principally as follows: Temples very distinctly convex; 
horns on a common base; sculpture throughout thorax denser and 
coarser; apophyses short and stout. 

Head and thorax black and yellow (see description of coloi- p:ittcrn, 
p. 141); mandibles entirely black; scape yellow below; annulus on 
flagellar joints (5) 6 12 (13) ; orbital ring entire; mesoscutum with a 
Finglc median yellf)w spot; scutellum entirely yellow; mesosternum 
yellow only along sternaulices; propodeal markings extending broadly 



166 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

forward from apophyses, but not reaching carina. Legs ferruginous, 
tibiae and tarsi yellowish ; front coxa yellow in front, black behind ; 
middle coxa yellow in front, piceous to ferruginous behind ; hind coxa 
ferruginous, with a large yellow spot above and a small piceous spot 
in apical middle ; wings yellowish, venation brownish ; abdomen black 
and yellow, petiole entirely black. 

Type locality. — Colima Volcano, Mexico. 

Type.— OwQ female in the U. S. National Museum (No. 57075), 
collected by L. Conrad. 

24. CRYPTANURA PROPINQUA (Cresscn), new combination 
Mesostenus propinquus Ceesson, Proc. Acad. Nat. Sci. Philadelphia, 1873, p. 152. 

Female. — Length 13-16 mm. 

Structurally very similar to aclnaces but with temples weakly con- 
vex; apophyses rather slender. 

Body color almost exactly as in aclnaces except that the mandibles 
apparently are always yellow at the base and the orbital rings are 
very narrow or even interrupted in the temples, and the scutellum 
is more or less black at the base ; but the legs are yellow, with the front 
and middle coxae, trochanters, and femora piceous behind, the hind 
coxa with a broad piceous stripe on the outer side of the upper sur- 
face and a shorter one on the inner side, the hind trochanter piceous 
with a trace of yellow on the upper and lower surfaces of the basal 
joint, and the hind femur piceous above and on the inner surface. 

Before me are a paratype and another female from Mexico (Fron- 
tera, Tabasco) ; a pair from San Jose, Costa Rica (M. Valerio, No. 
101) ; two females from Costa Rica (Paul Serre, 1920) ; and one female 
from Nicaragua (Mniszech, 1871), the last three received from the 
Paris Museum. 

The male has the malar sj^ace shorter, the frontal horns more slen- 
der and separated, the apophyses reduced to high elevations in the 
apical carina, which is distinct and sharply angulated medially, the 
petiolar area defined laterally by curved carinae, the apical slope of 
propodeum not distinctly transversely striate and the space between 
the carinae longitudinally striate, and the abdomen small and slender 
and subpolished. 

25. CRYPTANURA PRETIOSA (Viereck), new combination 

Polyacnidia pretiosa Viereck, Proc. U. S. Nat. Mus., vol. 46, p. 382, 1913. 

This species differs from all the other species with red abdomen 
which follow it in the present arrangement in its entirely red front 
and middle femora and in the form of the yellow marking of the 
lower division of the metapleuron. The latter is in the form of an oval 
spot extending forward from the hind coxa, whereas in the other 
species it either extends the entire length of the sclerite or has an 
angular forward extension above. 



ICHNEUiMON-FLIES OF GENUS CRYPTAXURA — CUSIIMAN 1G7 

The type and all()(yi)e are in tlie K("ini<;liclip Zooloj^jisclie !Nru?ouni, 
Berlin, the material examined in this study consist in«i; of a male para- 
type from Villa Mora, Parajjuay, and a female from Georgetown, 
British Guiana, identified by myself. The latter specimen agrees 
perfectly with the original tlescription. 

This is probably synonymous with spUonota (Cameron). 

26. CRYPTANURA GRACILIPES. new species 

Notable principally for its very slender legs. 

FernaJe. — Length 14 mm., antenna 12 mm., ovipositor sheath 5.5 mm. 

Temples distinctly concave; occipital carina high, especially at lower 
end, where it curves sharply mesad to join the hypostomal carina 
far back from the base of the mandible; vertex in front view slightly 
concave; frons with a few short striae radiating from ocelli; horns 
short and stout; eyes bulging, weakly convergent below; face some- 
what elevated medially, polished and very sparsely punctate, somewhat 
rugose above and in a shallow impression on each side of middle; 
clypeus polished and sparsely punctate, subnasute in profile; malar 
space nearly as long as basal width of mandible ; cheeks in front view 
nearly straight; antenna 35-jointed, slender, distinctly though not 
strongly thickened and flattened toward apex. Thorax rather unusally 
smooth, pronotum with humeral margin carinately angled anteriorly, 
upper portion, including humeral welt, polished and sparsely punc- 
tate, scrobe striate; mesoscutum polished, sparsely punctate, lobes 
weakly convex; scutellum narrow, convex, polished; mesopleuron 
])olisht'(l and punctate below, obliquely striate above; sternum and 
upper division of metapleuron polished and sparsely punctate; lower 
division of metapleuron coarsely obliquely rugosostriate; pro])odcuni 
basad of carina polish(Nl medially, rugose laterally, behind carina with 
striae radiating from between apophyses, posterior face transversely 
striate, bas;il median ai'ca defined, apophyses long and slender; legs 
verj' slender, hind femur about eight times as long as deep, more than 
three-fourths as long as tibia, reaching beyond apex of abdomen. 
Abdomen rather slender, subpolished; fiist tergite polished, j^etiole not 
depressed, postpetiole broader than long but not abruptly widened at 
spiracles, rather weakly convex; second tergite jdiout three-fourths 
longer than broad at base; ovipositor sheath distinctly shorter than 
abdomen beyond fii'st segment; ovipositor slendei- juid of unifunn 
depth to the distinctly subsagittate apex. 

Head and thorax black and yellow, abdomen largely ferruginous 
(.see description of color pattern, p. Ill) ; orbital ring broadly inter- 
rupted in temple and narrowly so in malar space; scape entirely black; 
anmilus on flugellar joints (5)0 10(12) ; mesoscutum with a single 
discal spot; .scutellum black basally; tegula with a broad stramineous 
and piceous margin; mesosternum yellow only along .sternaulices; 



168 PROCEEDINGS OF THE NATIONAL MUSEUM vol. m 

metapleuron broadly black along lower and anterior margins; pro- 
podeal markings barely including apophyses; front and middle legs 
yellow with femora and front coxa black behind, middle coxa ferru- 
ginous behind, hind leg ferruginous, the tibia and tarsus and a small 
spot at base of coxa above yellow, femur with a narrow dark streak 
above; wings hyaline, venation brownish; petiole yellow, postpetiole 
piceous with apical margin narrowly yellow. 

Male. — Like female but smaller and more slender; frontal horns 
longer and more slender; antenna 38- join ted; propodeum with 
apophyses short and thick, apical carina distinct and bent abruptly 
forward medially; abdomen much more slender, second tergite more 
than twice as long as broad at base. Annulus on flagellar joints 
(9)10-16(18) ; postpetiole only very narrowly yellow at apex. 

Type locality. — Trinidad Rio, Panama, 

Types. — Two of each sex, a holotype female, allotype male, and a 
male and a female paratype, all collected at the above locality by 
August Busck, March 17-30, 1912. They are in the U. S. National 
Museum (No. 57076). 

The paratypes are somewhat smaller than the type and allotype. 

27. CRYPTANURA GRACILIS, new species 

Female. — Length 14 mm., antenna 12 mm., ovipositor sheath 5 mm. 

Temples very weakly convex, more than half as broad as short diam- 
eter of eye ; occipital carina moderately high, curving slightly inward 
at lower extermity, in side view very nearly parallel to posterior 
margin of eye ; vertex straight across ; f rons with striae radiating from 
ocelli, horns short and thick, separated ; eyes weakly convergent below ; 
face polished, sparsely punctate, transversely striate above and in a 
shallow impression on each side of a low median elevation; clypeus 
polished, sparsely punctate, very strongly convex ; malar space nearly 
as long as basal width of mandible; cheek distinctly less than twice 
as broad as malar space; antenna 38-jointed, slender, barely thickened 
and weakly flattened toward apex, the joints in the thickened portion 
fully as long as thick. Thorax distinctly compressed ; humeral margin 
of pronotum transversely striate and with a strong carinate angle 
anteriorly, scrobe striate; mesoscutum coarsely and rather densely 
punctate, scutellum narrow; mesopleuron obliquely striate above, 
finely punctate below; sternum finely punctate; upper division of 
metapleuron punctate, lower division coarsely obliquely rugosostriate ; 
propodeum with median basal area defined, lateral areas largely rugose 
but polished and punctate in middle; area behind carina irregularly 
rugose, posterior face coarsely transversely striate, apophyses long 
and slender ; legs slender, hind femur fully six times as long as deep, 
and fully three-fourths as long as tibia, but not quite reaching apex of 
abdomen. Abdomen subopaque, finely shagreened beyond first tergite, 



ICHXEUMOX-ILIES OF GENUS CRYPTANUHA — CUSHMAN 169 

narrow, petiole not distinctly depressed, post petiole as lonp^ as broad, 
weakly convex; second tergite about three-fourths longer than broad 
at base, and hardly a half broader at apex than at base; sheath dis- 
tinctly shorter than abdomen beyond first segment, ovipositor slender, 
of nearly uniform depth to the subsagittate apex. 

Head and thorax black with yellow markings (see description of 
color pattern, p. 141) ; orbital ring interrupted in temple; clypeus yel- 
low only in middle, the black of the suture extending upward in the 
facial impressions; scape entirely black; annulus on flagellar joints 
(6) 7-19 (12) ; mesoscutum with a single median spot ; scutellum black 
in basal middle; mesosternum with a small trace of yellow next to 
sternaulices; metapleuron broadly black below and anteriorly; meta- 
pleuron posteriorly and metasternum ferruginous; propodeal mark- 
ings extending only a short distance before apophyses; front and 
middle legs yellow, black or piceous behind to apices of femora, middle 
coxa ferruginous behind, hind leg ferruginous with tibia and tarsus 
yellow; wings hyaline, venation blackish; abdomen entirely ferru- 
ginous. 

Type locality. — Villa Lutecin, near San Ignacio, Misiones, Argen- 
tina. 

Type. — One female in the U. S. National Museum (No. 57083) taken 
by E. R. Wagner in 1910. 

28. CRYPTANURA GENALIS, new species 

Female. — Length 13 mm., antennae 11.5 mm., ovipositor sheath 
5.5 nmi. 

Very similar to gracilis, differing ])i-incipally as follows: Occipital 
carina very high, in sitle view di\eigiiig below from postei'ior margin 
of eye; frontal horns on a low common base; eyes parallel; clypeus 
f)nly moderately convex; clieeks sli-ongly convex and fully twice as 
broad as malar space; antenna 32-jointed, stouter and more distinctly 
thickened toward apex, the joints in the ihicktned portion distinctly 
transverse; mesosculnm very sparsely punctate, prescntum not striate 
along notaulice.s; lobes not flatteneil; scutellum broader; lower divi- 
sion of metapleuron punctatorugose; i)roj)o(leuiii without a basal me- 
dian area, basal legion polished, sj)arsely punctate behind, more 
densely and coarsely j)unctate behind caiina, rugosopunctate laterally, 
posterior face closely transversely stii.ile. ;ij)ophyses very short and 
thiek; legs stouter, hind femur not nearly six times as long as deep 
aiifl barely two-thirds as long as tibia. Alxlonien stouter, postix'tiole 
much bioader than long, second tergitc barely n Imlf longer than broad 
at base; sheath as long as aixlomen beyond first tergito. 

Orbital ring not interrupted in temjile; black of clypeal suture not 
meeting in middle and not extending upward into facial impressions; 
anindns on flugellar joints (5) 0-11 (13) ; mesostei-num cons|)icuously 
yellow along sternaulices; propodeid maikings triangularly produced 



170 PROCEEDINGS OF THE NATIONAL MUSEUM vol.98 

anteriorly ; petiole tinged with yellowish and postpetiole with apical 
margin yellow. 

Type locality. — Venodio, Sinaloa, Mexico. 

Type. — One specimen in the U. S. National Museum (No. 57077) 
received from B. Preston Clark. 

29. CRYPTANURA VARIEGATA (Brnlle), new combination 

Mesostenus variegatus BruixI!;, Histoire naturelle des insectes, Hymenoptera, 
vol. 4, 1846, p. 277, pi. 40, fig. 3. 

Closely related to the two preceding species and differing from the 
above description of gracilis only as follows : Temple flat, hardly half 
as broad as short diameter of eye; clypeus only moderately convex; 
antenna 32-jointed, distinctly thickened and flattened toward apex, 
the joints there strongly transverse; scntellum rather broad and more 
weakly convex; propodenm without defined median basal area, lateral 
areas largely polished and sparsely punctate, rugose only laterally. 
Abdomen rather broad, postpetiole broader than long, second tergite 
more than a half broader at apex than at base. 

Clypeus black only apically and laterally, the black not extending 
upward into the facial impressions; metapleuron and sternum not 
at all red; annulus on flagellar joints (4) 5-10 (12) ; propodeal mark- 
ings extending triangularly nearly to basal carina. 

One female in the U. S. National Museum, taken in October 1922 by 
H. Lang at Kamakusa, British Guiana. 

30. CRYPTANURA PARANENSIS, new species 

Female. — Length 13 mm., antennae (tips broken off), ovipositor 
sheath 6 mm. 

Very closely related to variegata^ from which it is distinguishable 
by its distinctly longer ovipositor sheath and by the fact that the 
occipital carina diverges more strongly below from the posterior 
margin of the eye. From variegata it differs further as follows: 
Frontal horns on a common base ; thorax laterally less strongly sculp- 
tured, the striation of mesopleuron confined to the upper anterior 
portion, the punctation of the lower portion and of the sternum finer, 
and the lower anterior portion of metapleuron smooth and polished ; 
scutellum yellow only at apex and in basal angles; mesosternum en- 
tirely black; propodeal markings extending only very sightly for- 
ward from apophyses ; abdomen somewhat more slender. 

Type locality. — Peixe Boi, Para, Brazil. 

Type. — One female in the IT. S. National Museum (No. 57078) taken 
November 27, 1907, by Miss H. B. Merrill. 

What may be the male is represented by three specimens received 
from the Paris Museum and collected by Manger in Brazil. The labels 
also bear the figures 11-53. In these specimens the propodeum has 
a distinctly defined petiolar area from the anterior end of which the 
apical carina extends laterally and forward to the bases of the 



ICHNEUMON-FLIES OF GENUS CRYPTANUIL\ — CUSHMAN 171 

apf»i)hyses. The only complete aiitennii is 4;3-jointe(l with the aniuiliis 
on tla<xellar points (10) Il-IG (lb). The postpetiole is piceous, the 
niicldle coxae are very lar<>;ely ferruginous, while two of the 
sjiecimens have the mesosterninn largely yellowish. 

SI. CRYPTANURA TENUITEREBRATA, new species 

Fcinale. — Length 17 mm., antennae 15 mm., ovipositor sheath 9 mm. 

Conspicuous for its large size and long slender ovipositor, this 
species is, nevertheless, very closely allied to pai'anensis and vaHegata^ 
tlilFering from the former principally by the characters employed in 
the key. In structure and sculpture it agrees very well with variegata^ 
while in detail of color, especially of scutellum and propodeum, it is 
more like parancn-sis. The flagellum is somewhat stouter basally than 
in either of the other two species, the first joint of the flagellum being 
distinctly less than six times as long as thick, and the antenna is 
38-jointed, with the annulus embracing flagellar joints (5) 6-19 (14). 
The ovipositor sheath is nearly as long as the abdomen and the ovi- 
positor very slender, and of uniform depth to the distinctly subsagit- 
tate apex. 

Type locality. — Rurrenabaque River, Beni, Bolivia. 

Typef<. — Two females, the holotype captured in November 1921 
by "W. M. Mann on the Mulford Biological Exploration; the paratype 
fr<jm Sapucay, Paraguay. Oct<;ber 19, 1902. They are in the U. S. 
National Museum (No. 57079). 

32. CRYPTANURA INCERTA (Cresson), new combination 

Mcftostnuix incertus Cbesson, Proc. Acad. Nat. Sci. rbiladeliihia, 1873, p. 161, 
ft'iiiale. 

This and the species next following differ from the preceding six 
species in the much shorter and stouter hind femur, which is barely 
two-thirds as long as the tibia. The antenna in this group is some- 
what more slender and less strongly thickened and flattened toward 
the apex. Otherwise these s|X'cies are, in structure and color i)attern, 
very similar to the preceding group. 

Female. — Length 11 mm., antennae U mm., ovipositor sheath 4 mm. 

Temple flat, very strongly receding, distinctly less than half as 
broad as .short diameter of eye; vertex very weakly convex; frons with 
striae radiating from ocelli, horns very small, on a common base; eyes 
weakly convergent below; face ])olished and sparsely j)unctate, 
striate above and in longitiidinal imi)ressions; cly|)eus very strongly 
convex, polished and sparsely punctate; malar space nearly as long as 
basal width of niainlible: mueh more than half as long as cheek; an- 
tenna slender, weakly thickened and flattened toward apex, the joints 
in the thickened portion weakly transverse. Thorax distinctly com- 
pressed; humeral margin of pronotum transversely rugose and cari- 
nately prominent anteriorly, .scrobes striate; me.so.scutum with lobes 
flattened, polished, sparsely and coarsely punctate; scutellum narrow, 



172 PROCEEDINGS OF THE NATIONAL MUSEUM vol.98 

moderately convex, polished, with scattered punctures; mesopleuron 
striate above, polished and sparsely punctate below, as are also the 
sternum and the upper division of the metapleuron ; lower division of 
metapleuron coarsely, obliquely striatorugose ; propodeum with basal 
median area defined, lateral areas polished and coarsely, sparsely 
punctate to rugose, area behind carina irregularly rugose, posterior 
face transversely striate, apophyses rather long and slender. Abdo- 
men rather stout, minutely alutaceous subopaque, petiole not depressed, 
postpetiole broader than long; second tergite little more than a half 
longer than broad at base, its sides broadly divergent; ovipositor 
sheath hardly as long as abdomen beyond first segment, ovipositor 
slender, of uniform depth to the subsagittate apex. 

Head and thorax black and yellow (see description of color pattern, 
p. 141) ; mesoscutum with a single median spot; propodeal markings 
obliquely truncate shortly anterior to apophyses. Front and middle 
legs yellow, with coxae, trochanters, and femora black posteriorly; 
hind leg ferruginous with femur more or less piceous apically, tibia 
entirely and tarsus except the more or less blackish apex yellow. Wings 
slightly infumate, especially around apical margin. Abdomen ferru- 
ginous, usually with apex of tergite 1 narrowly yellow and postpetiole 
piceous, though sometimes entirely ferruginous. 

Of this species I have examined six females and one male as fol- 
lows: The National Museum female cotype from Mexico; a female 
and a male taken by August Busck, March 23 and 27, 1912, at Trinidad 
Rio, Panama; a female from Alhajuelo, Panama, April 7, 1911, A. 
Busck ; and two females from an altitude of 2,000-3,000 feet, "Pinches 
& Perene Vs," Peru, received from the Geographical Society of Lima. 

The Peruvian specimens have the white markings somewhat less 
extensive, with mandibles and sternum entirely black, and the first 
tergite entirely red. 

33. CRYPTANURA MACULIFRONS, new species 

Female. — Length 11.5 mm., antennae (gone), ovipositor sheath 4 
mm. 

Similar to incerta, from which it differs virtually only as follows: 
Mesopleuron obliquely striate over most of its surface; face white 
only medially and in orbits, mandibles entirely black; propodeum dor- 
sally, metapleuron below, and metasternum entirely reddish piceous to 
ferruginous; white markings of thorax as in incerta except that the 
mesosternum is entirely black; legs as in incerta except that the front 
coxa is very largely piceous and the middle coxa is entirely ferrugi- 
nous, as is also the abdomen. 

Type locality. — Misiones, "Env. de San Ignacio, Villa Lutecia," 
Argentina. 

Type.— One female in the U. S. National Museum (No. 57082) la- 
beled "E. R. Wagner, 1900." 



ICHNEUMOX-FLIES OF GENUS CRYPTANUIL\ — CUSHMAN 173 
KEY TO ALL SPECIES OF CRYPTANURA* 

This key is based partly on specimens and is partly compiled from 
characters gleaned from published descriptions. Of necessity it de- 
pends very largely on color characters and is purely artificial. 

1. Thorax red and yellow, rarely partly black above 2 

Thorax blaek and yellow, rarely propodeuiu and metasternum more 

or less reddish 9 

2. Head red and yellow ruficeps Cushman 

Head black and yellow 3 

3. Abdomen black and j-ellow 4 

Abdomen more or less, usually largely, red 5 

4. Mesoscutum immaculate black : mesopleuron red : 

Propodeum immaculate red albomarginata (SzC'pligeti) 

Propodeum with a black cruciform mark. 

albomarginata var. (Sz^pligeti) 

Mesoscutum red or piceous with a yellow median spot ; mesopleuron largely 

yellow piceothorax Cushman 

5. Mesoscutum with two small marlis discally 6 

Mesoscutum with a single median spot 8 

€. Mesopleuron white 7 

Mesopleuron red bipartita (Brulle) 

7. Apical carina not developed similis (Sz^pligeti) 

Apical carina developed medially dicostata Cushman 

8. Mesoscutum black variegata (Brull6) var. 4 (Sz«5pligeti) 

Meso.scutum red rufa Cushman 

9. Abdomen red 10 

Abdomen bla/k and yellow 36 

10. Scutellum conical 11 

Scutelium not conical 12 

11. Metapleuron subconically elevated tuberculata Cushman 

Metapleuron not elevated scutellaris (Sz^pligeti) 

12. Hind femur black above, red or yellow below 13 

Hind femur red, at n)ost black apically 15 

111 Hind femur yellow below lucida (Szc^pligeti) 

Hind femur red below 14 

11. Scutellum entirely yellow basimacula (Cameron) 

Scutellum yellow only at apex volcanica (Cameron) 

1'). Hind C'lXa black and white 16 

Hind coxa red 17 

10. Humeral margin of pronotum conically elevated on each side. 

politigaster Cushman 

Humeral margin of pronotum not conically elevated coxata Cushinan 

17. Proi)odeuni with a pale spot in each anterior angle 18 

Propodeum immaculate ba.sally 19 

I'trrohtris ( Szejiligef i ) , lioplcuris (Sz«'pligetl), and longipes (Sz^'pligeti) 
and its male variety run to 17 but no farther by the descriptions.] 



' Spool flc nnmoH whoHc authors nro Riven In parcnthrHos nrp here for the flfRt time nneS 
In conitilnnf Ion with titp kpiktIc nniiic Cryplniiurn, oxcfpt In tlio rnno of rrli/pn (CrrsHon) 
and npinnria (Unill*'-). Tlip nnnic-H Intnrntnria Ciinioron, mrxlrana ("roHHon. jirniilnqua 
CrcHson. nn<l vcrarpacin CnmfTon wpr« formorly uiifli-r MctoKtrnua. The re«t of the nnnica 
here transferred have formerly been under I'lilydinua. 



174 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 95 

IS. Propodeum with yellow spots basally uniformis Brues 

Propodeum with reddish spots basally rugosa Brull4 

19. Middle femur entirely black fusciventris (Cameron) 

Middle femur not entirely black 20 

20. Front femur and usually middle femur black or piceous behind or below 21 

Front and middle femora entirely red 32 

21. Propodeai markings each with a narrow, tonguelike forward extensioa 

before apophyses 1 22 

Propodeai markings truncate basally 23 

22. Postpetiole red ; upper margin of pronotum tuberculate on each side anteriorly. 

boliviensis Cushman 
Postpetiole piceous; upper margin of pronotum merely carinate anteriorly. 

isthmus Cushman 

23. Mesosternum white at least along sternaulices 24 

Mesosternum not at all white 2& 

24. Hind legs very slender, femur apparently nearly 8 times as long as deep; 

temples in dorsal view very sharply receding and slightly concave. 

gracilipes Cushman 
Hind legs stouter, femur apparently not more than 6 times as long as deep ; 

temples flat or weakly convex 25 

[variegata (Brulle) varieties, nigripes Brull6, and striata Brulle run 
to 24 but not farther by the descriptions.] 

25. Hind femur short, distinctly less than two-thirds as long as tibia ; ovipositor 

sheath barely twice as long as first abdominal segment incerta (Cresson) 

Hind femur at least two-thirds as long as tibia ; ovipositor sheath distinctly 
more than twice as long as first segment 2ft 

26. Cheeks fully twice as broad as malar space; temple rather broad, its angle 

with the longitudinal axis less than 45 degrees ; ovipositor sheath much 

shorter than abdomen : 27 

Cheeks much narrower ; temple narrower, its angle with the longitudinal axis 
moi'e than 45 degrees ; ovipositor sheath nearly or quite as long as 
abdomen 28 

27. Apophyses short conical ; lobes of mesoscutum strongly convex and sparsely 

punctate; postpetiole broader than long, margined with yellow, 

genalis Cushman 
Apophyses long; lobes of mesoscutum flattened and densely punctate; post- 
petiole as long as broad, not margined with yellow gracilis Cushman 

28. Hypostomal carina distad of occipital carina shorter than malar space ; ovi- 

positor slender and of nearly uniform depth except at apex. 

tenuiterebrata Cushman 
Hypostomal carina distad of occipital carina as long as malar space, very 
high ; ovipositor stout, much deeper near apex than at base. 

excalibur Cushman 

29. Head in dorsal view with temples concave and much less than half as long 

as short diameter of eye ; scutellum black medially paranensis Cushman 

Temples flat and nearly half as long as short diameter of eye ; scutellum 
entirely yellow , 30 

30. Middle coxa black and yellow ; face entirely yellow atripectus Cushman 

Middle coxa red ; face with a black stripe on each side of middle 31 

31. Ovipositor as long as abdomen and much deeper apically than basally. 

platyurus (Brulle) 
Ovipositor much shorter than abdomen and of unifoi'm depth. 

maculifrons Cushman 



ICHXEUMOX-FLIES OF GENUS CKYPTAXURA — CUSHMAX 175 

32. PropcMh'al spots with ton^io-shapcil oxtonsions nntoriorly_ vnriegata ( I?iull«5) 
I'ropiMk'al siKjts witliout such extoiisidiis anteriorly or, if extciuliiif; forward 

from apophyses, the extensions truncate or tapering 33 

33. Mesosternum hhick 34 

Mesostenunn at least partly yellow 35 

34. Posti>etiole black hyalina Brull6 

PostiK>tiole red conica Cushuian 

3-~». Probably synonymous spilonota (Cameron) 

pretiosa (Viereck) 

36. Propodeum with two yellow spots basally 37 

Propodeuni immaculate basally 42 

37. Me.^oscutum immaculate discally though sometimes with yellow marginal 

markings 38 

Mei^o.scutum with two small markings discally 41 

38. Mes().scutum immaculate ectypa (Cresson) 

Mesoscutum with yellow marginal markings 39 

39. Mesosfutum with yellow lines laterally opposite tegnlae. 

quadrimaculata Cushman 
Mesoscutum with cuneiform markings on anterior lateral margin 40 

40. Prescutum yellow on each side nitidiuscula (Cameron) 

Presi'utuin immaculate mediostrigosa Cushman 

41. Hind femur black posteriorly bicarinata Cushman 

Hind femur with a black line above championi (Cameron) 

42. Me.so.scutum with a .single median spot or immaculate, rarely (spinaria) the 

median spot flanked on each side by a small mark on inner margin of 

lateral lobe 43 

Mesoscutum with two discal marks 51 

•13. Meso.scutum immaculate ablata (Cresson) 

Me.so.scutum maculate 44 

41. Prop( ideal spots with tongue-shaped anterior extensions, which rarely become 

confluent across median line 45 

Propodeal markings without such extensions 49 

4'). Hind coxa red ( 9 ) or black with a yellow spot above ( $ ). 

spinaria (Brulle) 

Hin<l coxa yellow with black markings above 4l> 

4";. Hind femur red septentrionalis Cushman 

Hind femur black and yellow 47 

47. Hind fennir bhuk both dorsally and ventrally planiscutellata Cushman 

Hind femur black only dor.sally 48 

48. Hind femur with a narrow median yellow line dorsally. 

lineatifemur Cushman 
Hind femur entirely black dorsally veraepacis (Cameron) 

49. Hind coxa and femur red acinaces Cusliman 

Hind coxa and fenrur blark and yellow 50 

50. Mcsosterimm yellow; r»vip<».sltor sbrath shorter than abdomen. 

lamentaria (Cameron) 

Meso.Hternuin yellow only lalfrally and on each side (if iniddle [xistfriorly ; 

ovli)o.sllor sheath subeqiuil to abdomen In It-nglh propinqua (CresHon) 

51. Me.sopleuron with a large yellow s|HJt anteriorly and a small one p«isteriorly ; 

sternum, p4'tioIe, and hind femur entirely black apophysis Cushman 

Mesopleuron with n single large ol)li(|ue mark, Joined jMisleriorly to a yellow 
line on sternum; petiole partly yellow; hind fennir black only above 52 



176 PROCEEDINGS OF THE NATIONAL MUSEUM voi-. 9s 

52. Petiole black above, yellow below; hind femur with a median yellow line 

above orizabensis (Cameron) 

Petiole black below, yellow above; hind femur entirely black above 5S 

53. Mesosternum largely yellow ; propodeal spots broadly truncate before 

apophyses, the latter long mexicana (Cresson) 

Mesosternum yellow only along sternaulices ; propodeal spots narrowed and 
tonguelike before apophyses, the latter short bilineata Cushmaa 

ADDENDUM 
CREMNOCRYPTUS, new genus 

Genotype. — Polyaenus spiniferus Cameron. 

This genus will run in all existing keys to {Polyaenus Cresson) = 
Oryptanura Brulle, but the frontal horns are of quite different form, 
being flattened below and compressed above and separated by a deep 
groove. It also differs by the following characters : Ocelli situated on 
the sides of a distinct, sometimes very high, elevation; antenna in 
female only slightly thickened, slightly flattened on the outer upper 
side but not below; occipital carina strongly sinuate at lower ex- 
tremity; mandible distinctly tumid at upper basal angle; upper 
margins of pronotum tumid, but not angulated anteriorly by the 
epomia, which are weak or obsolete; notaulices shallow and extend- 
ing only about halfway to scutellum ; sternaulices short and shallow ; 
scutellum very broad, very weakly convex, and with sparse, coarse 
punctures; nervulus interstitial or very nearly so; abdomen coarsely 
and deeply punctate on basal three tergites, male abdomen fusiform; 
ovipositor neither distinctly subsagittate nor swordlike, but rather 
tapering and distinctly flattened above at apex. 

CREMNOCRYPTUS SPINIFERUS (Cameron), new combination 

Polyaenus spiniferus Cameeon, Journ. Straits Branch Roy. Asiatic Soc, No. 46, 
p. 117, 1906. 

Five specimens of each sex from Borneo, one female from Mindanao, 
and two females and one male from Singapore are before me. These 
agree exactly with Cameron's description. 

CREMNOCRYPTUS CINGULATUS (Tosquinet), new combination 

Polyaenus cingulatus Tosquinett, Mem. Soc. Ent. Belgique, vol. 10, p. 45, 1903. 

There can be no doubt that this New Guinea species is congeneric 
with the genotype. A considerable series, including both sexes, from 
Mindanao, Basilan, Samar, and Luzon belong, I suspect, to the species, 
though I hesitate definitely to identify them as such. They differ 
from spiniferus in having the stemmaticum strongly elevated above 
the ocelli ; the thorax, especially the posterior face of the propodeum, 
much more sparsely sculptured; the basal tergites more coarsely and 
less densely punctate; the antennal annulus constantly shorter; and 
from all but the Mindanao specimen in the blackish color of the entire 
inner side of the hind tibia. 

U. S. SOVERHMENT PRINTING OFFICE: 1948 



PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 



issued i'-f^jNoA, VJiJ^ l>y '^ 




SMITHSONIAN INSTITUTION 
U. S. NATIONAL MUSELIM 



Vol. % Wuhington: 1945 No. 3194 

NEOTROPICAL LANTERNFLIES OF THE GENUS PHRIC- 
TUS IN THE UNITED STATES NATIONAL MUSEUM, 
WITH DESCRIPTIONS OF FOUR NEW SPECIES 



F)V John S. Caldweix 



The Neotropical genus P/irlcfy^<i (Homoptcra: Fulgoridae^) was 
established by Spinola in 1839- for the unique species Fulgora dia- 
dema Linnaeus. Since that time various species have been described 
by Signoret, Distant, Schmidt, Lallomand, and Metcalf; in 1905 
Schmidt presented a key to the known species, which was modified 
by Metcalf in 1938. 

For such large and truly spectacular insects the species are relatively 
little known and their classification is in a confused state, probably 
because their descriptions have Ix^en extremely inadequate and very 
few have been illustrated. Specific identification has been based large- 
1\' on color and marking, characters that are often variable in intensity 
and exactness of pattern, wliile the structural characters generally 
emphasized concern the fmni of ihc giDtesque cephalic process, which 
in most cases cannot he adequately described. The length of this 
structure relative to the length of the pronotum has, in the past, 
served to segregate llu' species into gr(»Mi)s, which in tiiiii have been 
broken down into the respective species on the basis of color. This 
methofl (»f identification i-' almost a neressary evil, because the most 



' Kvon Hioiit'h Fulfjorn I.lnnnciih' jm n Hynonyni "f Lntrrnarin IJnnnoiiH (Fonnnh. Vrnr. 
Ulol. Sor. W.-iHhlni:tr«n. vol. .17, pp. 43-44, 1044), In the Hplfcllon of ii family nnmo I follow 
the* prlnrlplc proiuiHod by C. W. SnIiroHky (V<tIi. VII Hit. KonK. Knt., vol. 1. pp. fldU 0(13. 
1939). 

' Ann. Sor. Knf. Frnnre. vol. 8. pp. 218-221. 18.30. 

177 

«2M»r)7 1.') 



178 PROCEEDINGS OF THE NATIONAL MUSEUM vol.88 

pertinent structural characters of both the cephalic process and the 
genitalia defy description. The median notch in the caudal margin 
of the pronotum and the furcation of the median carina around this 
notch also furnish constant specific characters. In final analysis the 
male aedeagus, first illustrated by Metcalf, is of excellent value for 
identification purposes, but it must be remembered that the aedeagi 
in the Fulgoridae are inflatable and that they present a very different 
appearance when inflated than when deflated. The females possess 
good genital characters in that parts of the first valvulae are heavily 
sclerotized and ornate with spurs and ridges. Although these valvulae 
are not radically differentiated among the species, they are relatively 
constant wfthin each species, and a comparison of the accompanying 
drawings will readily demonstrate their specific value. 

The purpose of this paper is to present the more pertinent structiu'al 
characters of the genus, together with characters of color and pattern, 
in the hope that some of the present confusion may be cleared up and 
that recognition of the species may be made a much easier task. 

Because four undescribed species are added here, and because 
notatus Lallemand is not included in any existing key, a revision of 
the key is presented here. Unfortunately, xanthoptei^A Schmidt^ 
and notatus Lallemand * are known to the writer by their descriptions 
only. The order in which the species are discussed follows a tentative 
phylogenetic arrangement. 

ARTIFICIAL KEY TO THE KNOWN SPECIES OF PHRICTUS 

1. Cephalic process flattened apically, transversely arcuate, lacking definite 

apical teeth auromaculatus Distant 

Cephalic process with 5 apical teeth quinquepartitus Distant 

Cephalic process with 3 apical teeth 2 

2. Hind wings with large hyaline apical spots ocellatus Signoret 

Hind wings without hyaline apical spots, but small pruinose areas some- 
times present 3 

3. Elytra scarlet, continuous broad yellow transverse fasciae present. 

tripartitus Metcalf 
Elytra some shade of green or brown, transverse fasciae if present inter- 
rupted medianly 4 

4. Basal area of hind wing yellow, golden, or orange 5 

Basal area of hind wings red to scarlet 8 

5. Black or fuscous area in hind wings covering apical three-fourths ; cephalic 

process longer than pronotum regalis, new species 

Black or fuscous area in hind wings covering apical one-third or less ; cephalic 
process shorter than pronotum 6 

6. Basal area of hind wings golden yellow; elytra flecked with yellow and 

brown xanthopterus Schmidt 

Basal area of hind wings orange 7 



3Bnt. Zeit. Stettin, vol. 71. pp. 144-14R. 1910 (Ecuador). 
<Ent. Tidskr., vol. 52, pp. 188, 1931 (Ecuador). 



L.\NTERNFLIES OF GENUS PlIHICTUS — CALDWKLL 179 

7. Klyini jellowisli, irregularly Uiiisculale uver all with brdU n ; liiiul wings not 

uiaoulate sordidus, new species 

Elytra brown, inaeulute in basal two-tbirds with small round orange sjiots; 

hlud wings maculate with brown notatus Lallemaud 

8 (4). Elytni green in basal two-thirds, with a few small round red or orange 

maculae; apical third with large brown maculae 9 

Elytra brown or reddish brown 10 

It. Tet'tli in ajHi ot' cephalic process obtuse, somewhat deflected caudad ; median 

carina of pronotuni delinitely forke<l caudad inoebiusi Schmidt 

Teeth in aix'X slender, not dellected caudad; pronotal carina not delinitely 
furcate hoffmannsi Schmidt 

10. Elytra some shade of reddish brown, with pink calloused areas present, es- 

I)ecially basally; cephalic process toothed on ventral surface. 

diadema (Linnaeus) 
Elytra brown, maculate with y«'llow ; cephalic process smooth beneath__ 11 

11. Combined length of c«*phalic pincess and head as long as pronotum; expanse 

of trifurcate aiM'X equal to distance between (xular si)ines. 

minutacanthis, new species 
Combined lengrth of cephalic process and head longer than pronotum; ex- 
panse of apex much greater than distance between ocular spines. 

punctatus, new sp<H'ies 

PHRICTUS AUROMACULATUS Distant 

Pi-VTK 7. Kk.ukks C. L'4: F\j>.tf. S. Fu.ukks 1, IL' ; Plate 0; Pl.\te 10 

Phrk-tu.s uuromaruhitu.s I)isrANr, Aim. Mag. Nat. Hist., vt»l. Hi. ser. 7. pp. 072-673, 
l".m.'). 

'riieie may Ije .some doubt regiirdiiij^ the identity of this species, 
because Di.staiit makes no definite statenuMit as to whethef the apex 
of the ceplialic pi'ocess is trifurcate or flattened. He <loes mention tlie 
carinate aiileiior marojiii and states tliat the process is sliorter than in 
previously (h'.scfibed species. In the specimens studied not only is 
the pfocess veiy short liut also the apt'X is flattened and tiansverse. 
The cohjr of the hind wings is golden and not bi'ight yt'ih)w ; otherwise 
the description matches the specimens. Length oxer all, male .■*.."' mm., 
female oG mm.; elytra. male2G nun., female i'!) mm. 

Male bearing the data: "Tiimiiifasa, Bolivia. Dtc. Mill ford Biol. 
Expl.. I'.t'-M I'-ii'-J (W. M. Mann)"; female, "Ixiamas. Bolivia. .Miilfoid 
Biol. Expl., lU-2\ -I ( M. K. I>)pez)." 

'I'ype locality : Pxili via. 

I'MKK TIS OCKI.I.ATrS Siiinont 
I'l.ATi-: 7, Ki<ii iiKs I», 21 ; Pi_\tk S. FniUiiK W 
I'hrittuK urrlluluH SK.NoKi'rr, null. Soc. Km. i'niiice, vol. :',, ,ser. .3. p. v, 1S.'">5. 

As yet this is the only known .species with large hyaline areas williin 
the dark area of the hind w ings. The (•( plialic process is htnger than 
the pronotum. The caudal margin of the pronotum is dee[)ly not<he<l 



180 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

at the middle with the lateral margins of the notch sharply elevated 
and acute caudad. Length over all approximately 41 mm.; elytra 
32 mm. 

One badly damaged female from "Colom." [Colombia?], Baker col- 
lection. 

Type locality : Venezuela. 

PHRICTUS SORDIDUS. new species 

Plate 7, Figukes 3, 20 ; Plate 8, Figures 1, 13 ; Plate 9 

Allied to ocellatus in general pattern of elytra but lacking ocellate 
spots in the hind wings. Length, male 40 mm., female 46 mm. ; elytra, 
male 30 mm., female 37 mm. 

Dorsum of cephalic process, vertex, clypeus, and median stripe of 
pronotum light gray. Lateral and ventral margins of process dull 
cinnamon. Lateral areas of pronotum chocolate-brown. Elytra brown 
on basal two-thirds, irregularly maculate with dull yellow; broad 
transverse fascia present at base of reticulate area, broadly interrupted 
in center by a conspicuous brown dash. Apical third light yellow, 
maculate with large, irregular brown spots. Hind wings dull faded 
orange in basal two-thirds, apical third fumate. 

Cephalic process shorter than pronotum ; trif urcate apex with very 
obtuse teeth, especially laterally; median tooth somewhat deflexed 
caudad. Posterior margin of pronotum broadly notched; median 
carina ending at base of notch. First valvulae in female not trif urcate 
apically. 

Male holotype, U.S.N.M. No. 57224, Ecuador (Goodfellow), Goding 
collection. Female allotype, Quevedo, Ecuador (F. Campos R.), and 
one headless female, Ecuador, Goding collection. 

These specimens appear very faded, and it is unknown whether the 
color is natural or the result of immersion in preserving fluid. With 
regard to color they resemble the species figured and described by Met- 
calf as diadema; however, the cephalic process is distinctly shorter 
than the pronotum and therefore they cannot be that species. 

PHRICTUS MINUTACANTHIS. new species 

Plate 7, Figures 2, 16 ; Plate 8, Figure 2 ; Plate 9 

Resembles ocellatus in color and marking but differs from it in lack- 
ing the ocellate areas in the hind wings and having darker brown 
elytra. It is much smaller and more brightly colored than sordidus. 
Length, female 37 mm., elytra 30 mm. 

Entire venter, clypeus and face, lateral area of cephalic process, and 
dorsum of prothorax except median stripe dark brown. Dorsum of 
head, cephalic process, and median stripe of pronotum dark gray with 
apex of cephalic process rose to red. Elytra brown, fairly evenly 



LANTERN FLIES OF GENUS PHRICTUS — CALDWELL 181 

maculate with dull yellow, transverse fascia at base of reticulations 
irre<;ular and imperfect. Hind wings light red in basal two-thirds, 
apical third fuscous. 

Cephalic process slemler, shorter than pronotum; tritid apex very 
small, especially the median tooth. Caudal margin of pronotum with 
broad median notch, outer angles of notch abruptly acute and greatly 
elevated; median carina of pronotum apparently not forked around 
notch. First valvulae in female short, broad; apices more sharply 
angled than in oceUatus. 

Female holotype, U.S.N. M. No. 57225, fium Chaciuimayo, Peru, 
February 15, IDlS (C. H. T. Townsend). 

PHKICTUS PUNCTATUS, new upecieb 
Pi^TE 7, FiiiUBES 12, ii3 ; Platk 8, Figire 6; Pi-iTK I) 

Siuiilar to ininutacanthu but much larger and with more brightly 
colored elytra. The cephalic process is longer, with the apical teeth 
more acute. Length, female 44 mm., elytra 32 mm. 

Clypeus very light brown. Face and lateral margins of cephalic 
process cinnamon-brown. Vertex, dorsum of cephalic process, and 
median stripe on pronotum dull yellow. Elytra brown w^ith bright- 
yellow maculae groupt'tl toward the transverse fascia; claval area dull 
yellow; costal margin with two conspicuous yellow spots before 
leticulate area; transverse fascia bright yellow, narrow, interrupted in 
nnddle by a dark-brown dash more or less surrounded by yellow 
maculae. Maculae in right elytron tending to form an oblique strip 
U'tween the dark-brown ditsh and center of clavus. Hind wings 
hyaline-carmine in basal two-thirds, apical third light fuscous. 

Cephalic process equal in length to pronotum; trifid apex slender, 
with broader expan.se than width of head including eyes. Caudal 
margin of pronotum broadly concave, with a small notch in middle of 
concavity. Median carina definitely forked caudally around notch, 
with surface of pron(;tum on each side elevated above the forked 
carina. First valvulae in female with apices broadly concave on outer 
margins. 

F'emale holoty|)e, I'.S.N.M. No. 5722G, bearing the following data: 
"? IJugaba, Panama, collection Wm. Schaiis.'' One female paralype 
fiom El Volfdn, Panama, March 17, 11)43, is in the collection of the 
American .Museum of Natural Plistory. 

PHRICTUS HOFf'MANNSI Srhmidt 

1'i.An-; 7, FtoUKKH 10. 18; I'i.ait-: 8, Ji'k.ukk J> ; Tlatk 10 

I'hrictUH hoffmantiHi S< hmiht. Knt. Z<-1|. Sit-tiin, vol. iV,, pp. ,'{,'i8 .'140, l!M>r>. 

This s|)ecies most clearly resembles moebiu.si Schmidt but is dis- 
tinguisiusJ frouj it by a narrower cephalic process, with (he apical 



182 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

teeth slender and lying in the same place. It also resembles notatus 
Lallemand in having the elytra sometimes brownish and maculate, 
with small, round orange spots, but is separated from notatus by hav- 
ing the basal areas in the hind wings red and immaculate instead of 
orange and maculate with brown. 

Length, male 37 mm., female 46 mm. ; elytra, male 28 mm., female 
37 mm. Unfortunately, pests have destroyed the internal male 
genitalia. 

Represented in the collection by one male with no accompanying 
data other than "Goding Collection,-' one female from below Macas, 
Ecuador (E. W. Rorer) , and one female from Baiios, Ecuador, altitude 
1,800 meters (F. Campos R.). 

Type locality: Peru. 

PHRICTUS MOEBIUSI Schmidt 

Plate 7, Figures 11, 15 ; Plate 8, Figube 5 ; Plate 10 
Phrictus mofbiusi Schmidt, Ent. Zeit. Stettin, vol. 66, pp. 340-342, 1905. 

Resembles hoffmannsi in general appearance but differs in having 
apical teeth of the cephalic process more obtuse, with the median tooth 
deflexed caudad, elytra possibly lighter green, and with maculae fewer 
in number but brighter, and apices of first valvulae of female very 
acute. Hind wings differ from those of notatus in that the colored 
areas are red and scarcely maculate. 

One female bearing the following data : "Medellin, Vy. and Porce" 
(F. L. Gallego M.). Probable locality : Medellin, Colombia. 

Type locality : Colombia. 

PHRICTUS DIADEMA (Linnaeus) 

Plate 7, Figures 1, 4, 13. 14 : Pi^te 8, Ficiures 11, 14 ; Piatb 10 
Fulgora diaderna Linnaeus, Systeuia naturae, ed. 12, vol 1, p. 703, 1767. 

Although dkulenia is one of the more common species in the genus, 
its exact status is evidently still confusing. It was first figured by 
kStoU,^ but either the figure is erroneous or diaderrm is unknown today. 
The hind wing in the figure is unlike that of any known species, and 
the black color includes a much larger area than in typical diaderna. 
The figure by Drury ° (as annata) shows a much more typical pat- 
tern, while the photographic reproduction by Costa Lima ^ is dmderna. 
as accepted by most workers today. 

The general color of the elytra varies from light buff to dark shades 
of brown and sometimes even appears greenish ; however, in all this 



" Representation exaetement colorize d'apres nature des Cigales et des Punaises 
(Cigales), pi. 5, fig. 22, 1780. 

* Illustrations of natural history, vol. 3, pi. 50, fig. 4, 1782. 
' Insetos do Brasil, vol. 3, p. 45, fig. 40, 1942. 



LANTERNFLIES OK GENU8 IMllUCTUS CALDWELL 183 

variation pinkish oalloiisod areas aiv always |)n'S('Mt in the basal por- 
tions. The tiifuirato a|i(\\ of the ccpiialic i)roccss exhibits much 
variation as to size, deHeclion, antl form of the teeth. Tlie shape of 
the internal male fronitalia is constant and best shown by tlic illustra- 
tion (pi. s, tier. 14). The apices of the Hrst valvulae in the female are 
bluntly tiifurcate. Five females ranc^e in len<»;th from 4S to 52 mm. 
and four males from 41 to 46 mm. 

Records indicate that this spi'cies ranjjes throu<::h the (iiiianas into 
Brazil; also it has been recorded as a minor pest of cacao in Bahia, 
Brazil. In a letter to the writer, Pedrito Silva states that his reference 
to quinqiu'pnrfitiin Distant appearing in Tropical Agriculture^ is in 
error, as the pest on the cacao tree is diadenia (Linnaeus) and not 
quinqiu' part it ?/.*? . 

Type l(x.'ality: '"Indien" (probably Brazil). 

PHRICTUS REGALIS, new species 
Pl.\te 7, rn.UKKs 7. lit; Pi..\TE 8, F^atTRE 8; Piatf, 10 

Greatly resembling dwdema when the elytra are closed but with only 
the basal fourth of the liind wings yellow. Length M mm., elytra 36 
nmi. 

Median stripe on pronotum, vertex, and dorsum of cephalic process 
gray; apical teeth red. Venter of cephalic process brown, becoming 
fu.-cous towaivl clypeus; clypeus light yellowish. Elytra deep oliva- 
ceous, with calloused ai'eas and transverse fascia light red; costal and 
apical mai-gin lightly washed with black. Hind wings with basal 
fourth yellow, remainder black, with two or three large red spots 
present halfway to apex and three or four large yellow spots present 
in the black area neai" the basal fourth. 

Cephalic process stout, longer than pionotuni, l(Migtli and xcrtical 
height about equal; trifurcaic apex with I(»ng acute teeth. Caudal 
inai-gin of pidiiotnm scarcely in<lentc<l medianly; median carina 
appearing deeply bifiwcate. (Unfoi'tunately. most of this ai-ea is 
obliterated by a large pinhole.) 'J'rifurcale ajjices of Hrst female val- 
vulae with apical aiul outer teeth acute, inner teeth bliinl in latei-al 
aspect . 

Female holotype. U.S.X.M. No. r)7227, from Maroni Kiver. Ficnch 
(iiiiaiia. vicinity of I)Ms('i-rc ((i. Molicrg). 

IMIKKTUS UriN(JUKPAUTITUS Dintnnl 
I'l.AIT. 7, l-'KifRKS K, 17; I*T,ATK K, F-'KilKKS 10, ].'»; ri.AIK'.l 

I'hii'iuK iiuintfiu p'lrliluM Disiant. Itiuldiriji < 'ctil raii-AiiM'ricatia. IIi)innptrra. vnl. 1. 
P 2t. pi t. flir. 8. 18.S:?. 

This unii'^ua! '-pecies closely resembles triparlitux Mel calf in color 
and marking but is ilistinguisherl from ii bv the apex df the cej)halic 



•Trop, Acr. vol. 21. p. 12, 1044. 



184 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

process bearing five teeth. In the 15 specimens examined the apical 
teeth vary from acute to obtuse, and sometimes the intermediate teeth 
are much reduced in size. The longitudinal veins in the elytra are not 
so green as in Distant's illustration, and the bluish-white pruinose 
spots in the dark apical area on the hind wings are sometimes absent. 
In addition, the red areas in the hind wings are often less maculate 
apically with black. The shape of the internal male genitalia appears 
to be veiy close to that figured by Metcalf for tripartitus ; however, 
the expansion of the inflatable sacs is much less, and the writer be- 
lieves that these two species are distinct. As in diadema the two sexes 
vary considerably in length, the males ranging between 43 and 45 mm. 
and the females between 47 and 49 mm. 

All specimens are from Panama and the Canal Zone. 

Type locality : Panama. 

PHRICTUS TRIPARTITUS Metcalf 

Plate 7, Figures 5, 22 ; Plate 8, Figxjre 7 ; Plate 9 

Phrictiis tripartitus Metcalf, Bull. Mus. Comp. Zool., vol. 82, p. 365, pis. 20, 21, 
1938. 

Metcalf believes that this species may be the unnamed variety of 
diade/ma described by Walker ^ and figured in the Biologia.^" In 
general color, pattern, and size it approximates specimens of male 
quinquepartitus in the collection. Although the aedeagus of the male 
is of tlie same general form in both species, it is much more inflatable 
in quinquepartitiis ; in the female there are differences in the first 
valvulae. 

In the specimen believed to be tripartitus the teeth in the trifurcate 
apex of the cephalic process are very obtuse but between the median 
and right-hand tooth is a definite bump that may be construed as a 
vCvStigial tooth; on the other side of the median tooth the margin is 
crenulate. This specimen, measuring 51 mm., is longer than any speci- 
men of quinquepaTtitus. The difference in total length is probably 
accounted for by the longer cephalic process possessed by tripartitus. 

Female plesiotype from Virginia, Guatemala, November 1915 (Wm. 
Schaus) . 

Type locality : British Honduras. 



" List of homopterous insects in tlie collection of the British Museum, p. 264, 1851. 
'" Biologia Centrali-Americana, Honioptera, vol. 1, pi. 4, fig. 5, 1883. 



U. S. GOVERNMENT PRINTING OFFICE. 1945 



U S NATIONAL MUSEUM 



PROCEEDINGS. VOL. 96 PLATE 7 




CEPHALIC PROCESS IN PHRICTUS. 

Frontal profile of riylil half: 1. diaJema (fcmaU-); 2, minularanllns (male); 3, jorduhu 
(male); 4, diadrma (male); 5, triparlitus (entire profile, female); (u auromacualus ( <;niaU-): 
7 rfgalis (female); 8, quinquepartitw: (male); 9. nrrllalus (female, imperfect); lO.V'o/- 
mflnn/i (female); 11, mo/-tiM.rt (female) 12, punctalui (female) , ^ ,^ * • 

Lateral profile: 13, diadniu, fmale); 14, diadrmu (female); Is, ».oW;;»u (female); 16 m,«u/fl- 
canlhis (male); 17, quinqiupartilus (male); IS, huffmannn (male); P. ra'fl/i.f (f<-maie); 
20, jnrdidus (female); 21, ncfllaius (female, imperfect); 22, irtpartilui (female); 23, puncta- 
lus (female); 24, auromarulnius (fenjale). . 

Owing to their larv'c si/.e, rrRali^, diadnna, nui'u,nrt>,ir!ttu<. and iriparlUus are drawn approx- 
imately one-half scale. 



U. S. NATIONAL MUSEUM 



PROCEEDINGS. VOL. 96 PLATE 




Valvula and Genitalia in Phrictus. 

Lateral and ventral profile of first female valvula (left): 1, sordidus; 2, minutacanlhis; 3, 
ocellatus; 4. auromaculatus; 5, moebiusi; 6, punctatus; 7, tripartitus; 8, regalis; 9, hoffmannsi: 
10, quinqueparlitus; 11, diadema. 

Ventral aspect of male aedeagus: 12, auromaculatus; 13, sordidus; 14, diadema; 15, quinque- 
parlitus. 



U S NATIONAL MUSEUM 



PROCEEDINGS. VOL. 96 PLATE 9 





AUROhlACULATUS 



PUNCTATUS 





MINUTACANTHI S 



SORDIDUS 





^- •:* V 



■^ 



TRI PARTITUS 



QUINQUEPARTITUS 



SPECIES OF PHRICTUS DORSAL ASPECT 
auromaculalus, female; punclalus, female lioldtypc; minulacanthuf, female licl-nyiH- 
sordiduj, female allcjtype; Iripartitus, female; <iuinqufparlUu<, male. 



U. S. NATIONAL MUSEUM 



PROCEEDINGS, VOL. 96 PLATE 10 



c 





^ 



•\ 



REGAL/ S 



DIADEM A 








HOFFMANNSI 



MOEBIUSI 





AUROMACULATUS 



DIADEM A 



SPECIES OF PHRICTUS: DORSAL ASPECT. 

egalis, female holotvpe; diadema, female; hofmannsi, female; moebiiisi, female; auro- 
maculatus, male; diadema, male. 



PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 




issited 5' J5(V '^■^ 0?'l^i by the 
SMITHSONIAN INSTITUTION 

L'. S. NATIONAL MUSEUM 



Vol. % 



Wuhington : I94S 



No. 3195 



HYPOKIIAMPIirs l^VTRIS. A NEW SPECIES OF HEMI- 
KAMPIIID FISH FUO^r SIXALOA. MEXICO, WITH AN 
AXALYSIS OF THE OEXEKIC CHARACTERS OF HYPO- 
RHAMPH US AND HEMIRAMPHUS 



Hv RoilIIfT R. MlI.LKR 



Till. |i.iii<iiy of oui- kiiowU'tl^^c of tlu' fresh-Av titer fish fauna of 
MortliWfstiTii M('.\i<-o is cNident from the novelties which Ralph G. 
Milh-r has c'<tnected in that re*;ion in recent years. In addition to 
the distinctive I)oroi<omn fon'ithl Hubbs and Miller (19-H) and a new 
(riht being described by nie in Copeia, a new species of hallbeak of tlie 
;zenus IlyporhamphuJi is now made known. 

About GO years a^^o Meek and Goss (1885, p. 221) wrote that the 
American halfbt-aks rcfeired to /// mh-t/mp/n/s^ were "in a condition 
of gieat confusion."' Althoii^'h a niniibci- of papci's dcalin<^ with tluf 
New World .-pecics have appeared since liiat time, the systemat ic status 
and particnlaily the distriitut ion of the American forms are still far 
from clear. 

The discovery of the new halfbeak, described below, biin^^s nj) the* 
(juesiiou of the <reneric validity of //i/por/ttt//i/>/i>/s and has prompted 
u ciitical study of hifisilirtis'/s and un'ifaxciaiiis, the ^feiiotypes, I'espec- 
tively, of //rri4iromp/iu.s Cuvier and IIi/p(>rhain]>JiUi< (lill. This study 
has j)roved to be most pr"odiict ive. for a number of t rencliant and easily 
observed cliaraiters, lieretofnrr appaicntlv o\erlooked, weic found. 
The presence or ab-ejice of scales on the upjiei- jaw also was noted by 
Smith ( ll).'i:5, p. VM)). In |)reparin;j; table 1, in which the «^enotype.s 
of IlrmiramphuH and Uyporhamplivx are compared, I examined 135 
s|)ecimens of ^inifayciatns and 05 specimens of lir<i-'<irnnxis in the col- 
lections of the W S. Xational .MuH-um. These specimens represent 

' SiK-Urd llrmirhnmphun liy tlirni niiil liy i\ lioHt of otluT nuttHirH. Tlio rtrlk'iniil HpoMltifc 
bjr Cuvlor (1817. p. \HC) Ih Himi Unrnphun. 



r.n.'ir.ao— 4.'» 



IK.--, 



186 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

material from the known American range of both species : ■mufasciatus, 
from Cape Cod to Uruguay in the Atlantic and from San Diego - 
to Peru in the Pacific ; and hrasiliensis, from New York to Brazil. 

Gill (1859, p. 131) based Hyporhamphus principally on the tricuspid 
teeth (whence the name of the type species, H. trieuspidatus, a syno- 
nym of unifasciatus) , but he later (1863) found, and Poey (1860, 
p. 298) previously had noted, that Eemiramphus likewise has tricus- 
pid teeth. Poey's and Gill's observations on the nature of the teeth 
were correct, and hence I do not agree with Weed (1933, pp. 47, 57) 
and others who stated that the teeth are simple in Hemiramphus. As 
Smith (1933) has shown, and as I have also observed, the form and ar- 
rangement of the teeth vary with age and with different species. The 
jaws of a single individual may have unicuspid, bicuspid, and tri- 
cuspid teeth, and, in at least one American species, Hyporhamphus 
rosae (Jordan and Gilbert), only the largest individuals appear to 
have tricuspid teeth — hence the frequent statement that H. rosae has 
only unicuspid teeth. 

The fundamental characters distinguishing the American species of 
Heiniramphus and Hyporhamphus^ such as the presence or absence 
of scales on the upper jaw, the presence or absence of a bony rim along 
the side of the nasal fossa, and the arrangement of the sensory canal 
and pores on the preorbital (fig. 9), may be features that will separate 
world half beaks of this type. This is suggested to me by Smith (1933, 
pp. 130-131), who made a primary division in his key on the basis of 
a naked versus a scaled upper jaw, and b}^ the very few Old World 
half beaks I have examined. In Euleptorhamphus Gill, however, the 
upper jaw is scaled as in Hyporha7nj)hus, whereas the rim and the 
form of the nasal fossa and the sensory canal of the preorbital are 
essentially as in Hemiramphus. 

The pattern of the scales on the uj^per jaw, the shape of the pre- 
orbital, and the arrangement of the teeth maj^ be found to have generic 
or only specific value. The solution of these problems will necessitate 
a comprehensive review of the halfbeaks of the world. 

The form of the sensory canal and the pore on the preorbital are 
usually visible in Hyporhamphus^ but the overlying scales and skin 
must be dissected from this bone in Heniiramphus before the canal 
and pores can be clearly seen. The two pores shown near the upper 
end of the posterior margin of the preopercle in Hyporhamphus (fig. 
9, A) are apparently absent in Hemiramphus^ but this character was 
checked only on a comparatively few individuals of each genus. 

In table 1 I have abandoned the "key" characters — air bladder cel- 
lular or simple, sides of body vertical or convex, position and shape of 

2 In material from San Diego, Calif. (Stanford Nat. Hist. Mua. No. 9912) I found one 
specimen of this species, Which, to my linowledge, represents a northward extension of 
known range on the Paciflc coast. 



A NEW IlLMlHAMrillD IISIL — MlLLEIi 



187 



dorsal lin, ami position of pel vies — used by many writers to separate 
Ueininunjihu.s from Ilf/porfunnphiu^. The nature of the air bladder 
is diflicult to discern but may be of considerable phylogenetic impor- 
tance; the form of the sides of the body is an untrustworthy character 
because it is frequently rendered impractical by preservation; the posi- 
tion and shape of the dorsjil lin is not so distinctive a feature as is the 
ditVerence in the basal lenjiths of the dorsal anil anal fins; and the 
position of the pelvic fins is useful largely for specific or subspecific 
separations. 

Table 1. — Diaguustic differences bettceen Hyporhamphus unifaBciatue and Hemi- 

ramphus brasiliensis ' 



Character 




unifasciaiui 


brasiUensis 


Upper Jaw 


Sc-altti 


Naked. 


MaTKin of nasal 


fossa 


Surmounted by a prominent bony 


Lacking a bony rim in this position. 


(see flg. 9). 






rim along posterolateral border. 




Sensory canal 


on 


pre- 


Unbranched; with an exposed pore on 


Branched; witli a jiore at end of pos 


orbital (see flg. 9) 




side and another pore at terminus 


terior branch (wliich terminates in a 








of canal near anterior margin of 


bony rid^'c near front of orbit) and a 








nasal fofxsa. 


pore at end of anterior branch near 
antcroventral nisiri^'in of nasal fossa. 


I)'>r««l fiT! 






Over or nearly over origin of anal, its 
base and that of anal equal or sub- 
equal. 


In advance of anal oripin, its base 1.5 
to 2.1 times that of anal fln (1.3 or 1.4 
in young). 


Caudal Hu 






Moderately forked, the distance be- 
tween caudal ba.se and shortest 


Deeply forfceii, the dLsfancc between 








caudal base and shortest caudal rays 








caudal rays 7.-1 to 9.0 in standard 


12.5 to IC.3 in standard length. 








length. 




Sh:'.]!*.- cjf iiasiil 


fossa in 


Broad, and little dcpressc*!. Its great- 


Narrow and greatly depresised, Its 


a'JuIt. 






est inner diameter more tlian one- 
half that of orbit. 


greatest dianietor oni-fourth to one- 
third that of orbit. 



> Characters of Ilvjwrhamphvi confirmed on the type specimen fll.P.N.At. No. 8407) of the genus, H 
trtciupid(Uus(,=-unifafdiiliu), and of Jhniiramphun on "lopotyie.s" (sjieeimens from Jamaica, U.S.N.M. 
No. 30077) of //. briuUientiM. 

The new species described below is the lirst to be definitely recorded 
from fresh vater in the New \\'oild. It apjieais to be restricted to 
a fluviatile habitat, for a numbei- of collections of halfl)eaks along the 
west coast of Mexico in the n-gioii where the new species was dis- 
cov«'red ct^ntain no sj)ecies identical with it. 

I name this distinctive fish />'/// /.v,gcii it i\e of jxifrr (father), because 
my fathej-, lljilj)h (i. Miller, collected the It types hihI only known 
spCH.imeii.s. 

HVrOIUIAMI'HUS rATKIS, new iprciri 
I'LATK 11 

Tyin'8. — The holotype (U.S.N.M. N'o. 129950) is a mature aihilt 
(presumably a female, sec below), lis iiim. in standard length, and 
was collected on May 4, 1912, by IJalpii (J. Miller in Rfo del Fuerte, 
one-half mile above the town (»f Kl Fuerte, which is about 2(1 miles 
northeast of San lilas, Sinola, Mexico. Tlie 1.'5 paratypes (U.S.N.M. 



188 PROCEEDINGS OF THE NATIONAL MUSEUM vol. m 

No. 129957), 107 to 130 mm, long, wore collected with the holotype. 
One fish in the lot, a specimen 118 mm. in standard length, is tlie only 
individual of the series that has distinctly larger pectoral and pelvic 
fins. On examination it was found to be a ripe male. One of the 
others, a specimen 109 nun. long with short pectorals and pelvics, was 
found to contain eggs in various stages of development, some of them 
apparently fully mature. The remainder are ])resumably all females. 

Diagnosis. — A Hyporhaiiypku^ with pelvic fins about equidistant 
between caudal base and gill opening, 21 to 24 gill rakers on lower 
limb of first arch, with a relatively long mandible (3.6 to 4.2 in 
standard length), without scales on dorsal or anal fins, and without the 
fleshy tip of the mandible red. 

Desonption. — Bodj^ rather slender, its depth 8.0 to 9.6 in standai'd 
length, little compressed, the sides rounded; w-idth of body in depth 
1.05 to 1.4; head 4.5 to 5.0 in standard length; mandible (measured 
from tip of upper jaw to end of bony tip) 3.6 to 4.2 in standard 
length and 0.7 to 0.9 in head length (broken in one specimen) ; snout 

2.8 to 2,9 in head ; orbit 4.0 to 4.3 in head, 1.35 to 1.45 in snout, and 1.65 
to 1.85 in postorbital; interorbital 3.8 to 4.1 in head and 1.55 to 1.7 
in postorbital; length of preorbital 1.5 to 1.65 in orbit; depth of pre- 
orbital 1.5 to 1,75 in orbit; width of nasal fossa 1.85 to 2.15 in orbit; 
base of anal fin 1.01 to 1.08 in base of dorsal fin; pectoral short, 8.4 to 
9.35 in standard length in females (7.9 in the male) and 1.75 to 1.95 
in head (1.65 in male) ; pelvic 2.7 to 3.0 in head in females (2.25 in 
male) ; midcaudal rays (measured from midbase of caudal fin to tip 
of shortest middle ray or rays) 8.4 to 9.3 in standard length, 1.7 to 

1.9 in head, and 2.1 to 2.4 times the length of the orbit. 

The fin rays vary in number as follows: Dorsal 13 to 15, usually 14; 
anal 15 or 16. usually 16; pectorals 10-10, 10-11, or 11-11, ahnost al- 
ways 10-10; pelvics always 6-6. I depart from my usual method in 
counting the rays of the dorsal and anal fins and regard every element 
as a separate ray, because this procedure has been followed by virtually 
all students of this group of fishes. Without exception the first two 
rays of the doi*sal fin are unbranched, and the first two rays of the 
anal fin are also simple except in two specimens in which the first 
three rays are unbranched. 

The gill rakers on the lower limb of the first gill arch (counted on 
both sides) vary from 21 to 24. 

The lateral series scales (counted from upper angle of gill opening 
to caudal base) number about 53 to 59, usually 55 to 57; an accurate 
count is difficult to obtain because the scales are largely missing from 
the sides. 

The pelvic fins lie about equidistant between the base of the caudal 
fin and the gill opening, varying between the pectoral base and the 
middle of the opercle. The dorsal fin varies in position from equi- 



A NEW HEMIRAMPPIID FISH — MILLER 



189 



distant bctwi-en caudal base and pelvic insertions ((. niuch nearer 
{lelvic insertions tlian caudal base. 

The tei'th of the holotype are unicuspid, bicuspid, and tricuspid and 
are arranged in about three to seven irre<!;ular rows in the upper jaw 
and two to five rows in the lower jaAv. Tricuspid teeth are present 
only posteriorly in each jaw and virtually all the anterior teeth (from 
about the middle of each jaw forward) are unicuspid. In the region 
where unicuspid and tricuspid teeth intergrade, occasional bicuspid 
teeth occur. The tooth rows are conspicuously broader medially on 
each side of the upper jaw tlian they are at either end, and teeth are 





Figure 9. — Sketch of head regions of Ilyporhamphus and Ifemiramphus to illustrate 
certain diagnostic differences (see tabic 1): A, I/yporhamphui unifasciatus, 183 mm. in 
standard length, from Key West, Florida (U.S.N.M. No. 34599); B, Hemiramphui 
brasilieitsis, \h2 mm. long, from Key West, Florida (U.S.N.M. No. 38684). Drawn by 
•Mrs. A. M. Awl, U. S. National Museum. 

absent at the tips of both upper and lower jaws. In the lower jaw the 
rows of teeth are of nearly uniform width but are somewhat broader 
close to the proximal end on each side and theji become narrow gradu- 
ally forward and abruptly behind this region. In the largest ])ara- 
ty|>e (KiO mm. in standard length) there are more ti'icuspid teeth than 
in the holotype (118 mm. long), which agrees with my observation in 
//. roiiae that tricuspid teeth api)ear with increasing size of the indi- 
\idual (this was also not<'d in other American //rnnramj>hus and 
Ih/lforJicinijihuR). Otherwise the teeth of the paratypes have es.sen- 
tially the same form and arrangement as in the holotype. 

The triangular up{)er jaw is rather bluntly pointed at the apex and 
l>road<'r at the base than it is long, \\nien the mouth is closed most of 
the outer teeth of the lower jaw are exposed. The scales of the upper 
jaw are irregularly arranged, the transverse rows munbering five or 
six arross the base, tlion al»oiit four, whereas from about the middh' to 
tlio tip of the jaw thoy are bi^erial — with a single scale on each side of 



190 PROCEEDINGS OF THE NATIONAL MUSEUM vol. o« 

the slight median ridge. Although the scales cross this low ridge pos- 
teriorly they do not usually do so anteriorly. 

No scales were observed at the tip of the upper jaw, but these may 
have dropped off. The sides of the head, including the region of the 
mandible below the jaws, are covered with deciduous scales. 

The margins of the prolonged mandible or "beak" are nearly par- 
allel throughout, diverging little until the posterior end is reached. 
The nasal flap is small. 

The dorsal and anal fins are low, highest anteriorly, with rays 3 to 
5 longest ; these rays in the anal fin are almost three times as long as 
the last ray, whereas in the dorsal fin the anterior rays are only about 
twice as long as the last ray, which is slightly prolonged and falls 
some distance short of reaching the bases of the procurrent caudal 
rays. The asymmetrical caudal fin is very weakly forked, less so 
tlian in any other American species I have seen except H. rosae. As 
in many halfbeaks, the lower caudal lobe is longer. 

The air or swim bladder as noted in the single male is simple, with- 
out any cellular structure. 

Coloration. — The general coloration was noted in the field by the 
collector. When taken from the water the body of the new species 
was intense blue and green varying in brilliance according to the re- 
flection of light from the surface, the blue and green grading into 
each other. The fins or the belly are believed to have been yellow or 
orange. No bright color was seen anywhere on the beak. This ob- 
servation is important, for most, if not all, of the American halfbeaks 
have the fleshy tip of the mandible red. According to Herre (1944, 
p. 9) the Philippine species of Hemiramphus (including Hyporham- 
■pJius) have this tip red, green, or greenish white, depending upon the 
species. I therefore interpret the lack of red color on this structure 
in patris as a character of specific value. 

The color of the preserved specimens (in alcohol) is mostly light 
silvery to pale brownish.^ The back, above the lateral band on each 
side, is marked with brownish punctulations. which are usually more 
concentrated on the posterior borders of the scales. Along the middle 
of the back are three narrow longitudinal rows of dark pigment, 
broader near the occiput and particularly over the caudal })eduncle ; 
the outer rows are more or less continuous past the base of the dorsal 
but the middle row is disrupted in this region into a series of U- oi- 
V-shaped markings between the bases of the rays. The base of the 
anal fin is marked similarly to that of the dorsal base, but the longi- 
tudinal rows of pigment are far less conspicuous. On each side of 
the body is a dark band, probably silvery in life, which is very narrow 
anteriorly and broadest between dorsal and anal fins. The upper 
surface of the anterior part of the head, including the upper jaw, and 
of the mandible is black; the lower surface of the mandible is finely 



A .\K\V HEMIRAMPHII) Fli:H — MILLEK 191 

pi<;iiii'iit<'il with black cliroiiuitophorcs ftulinj^ po.steiiorly bo that 
both cliiii and tluuat art" hugely coh>rlc>;s. The tips of tlie caudal rays 
and those of iho Ioii«i:er dorsal rays arc marked with fine black jiunclu- 
lations; tiie other fins are mostly pale. Alon<i: the underside of tho 
caudal peduncle are three rather irregular lontritudinal rows of dark 
l>iLrinent. The silvery peritoneuni is ovei-lain by foppi'iy Ijiowii and 
by fine, black punctulations. 

Habitat and associates. — Rfo del Fuerte, near El Fuerte, Sinaloa, 
is a deep river with sand and mud bottom and abrupt rocky banks. 
On -May 4, r.»42, when the types were collected, the current was fairly 
swift, and hauls with a 2o-foot bag seine were made in water generally 
1 to .") feet deej) but more than G feet in places. At noon the air was 
37*^ C. and the water 32 C No vegetation was seen, and the shore 
was sandy, with trees along the bank. Collecting was confined largely 
to the backwaters. The point wlierc the fish were secured is fully 
KK) miles upstream from the Pacific. 

In the large colle«'tion made here, the following fishes, tentatively 
identified, wer«' also seined: A species of cyprinid fish of the genus 
Gila; two specimens of a catfish of the gemis Ictaluius; cyprinodont 
fi.shes of several genera including MoUienisia sphenops; six mullets, 
Af/onostonuis montieola: a large niunber of the fresh-water atherine 
Mclaniris cryataUinus ; and two gobies. Ait'aou,s (or Clumopkonis) 
transandeanus and Gohiomonis 7>wntlatv-s\ Most of these species are 
confined to fresh water. 

Ranfi< . — The new s[)ecies was collected only in the Kio del Fuerte. 
Ralph G. Miller -aw halfbeaks in the Rfo Culiacan at Culiacan, Sina- 
loa. Mexico, alxiiit l.">0 miles south of El Fuerte and about 40 miles 
ui)Stream fr(»m tin* Pacific, but the identity of this species is unknown. 

Relationships. — Tlyporhamphus patris appears to be the southern 
repi-esentative of //. ros,ic (.Jordan and Gilbert) (1880). which is 
known frojii San Pedro, Calif., south to the tip of Paja California, 
then up the west side of the Gulf of California and southward along 
(lie maiidand of Mexico to Guaymas. Sonora (Evermann and Jenkins, 
1801, p. 13.'); record confirmed by examination of the five specimens 
from (iuaymas in the Stanford Natural History M\iseum, No. 437). 
Ki'f) del Fiier-te, tlie liabital of jxitris, is about 170 miles soiilli of 
tiuaynias. 

'ilie two s|tecie^ agree in most measuremenl> and coimts and in the 
following important charact<'rs: (1) P(»st«'rior position of the pelvic 
fins; {'!) gill rakers: 21 to 2;') on the lower limb of the first gill arch 
in my counts for rosne^ 21 to 21 for patris; {W) long mandible, which 
api)ears to Ix' slightly longer in roKar^ but a series of comparable sizes 
would probably eliminate this difference; (4) no scales on i\\*.' dorsal 
or anal fins; (.'>) dentition. The tw(> species differ as slutwn in the 
(oiiiparison presente<j in table 2. Soine oi- all of these difrerenc<»s 



192 



PROCEEDINGS OF THE NATIONAL MUSEUM 



may vanish wlien larger series of both species from more localities 
are available, but it seems best at this time to regard them as distinct 
species. 

Table 2. — Comparison of Hyporhamphus patris and H. rosae 



Character 


Tosaf 


patria 


Color of mandible 


Dark red 


Black. 


Fleshy tip of mandible 


Red 


Not red. 


Scales in lateral series 


58-65 


54-59. 


Bony interorbital into length of 

head. 
Diameter of orbit Into head 


3.4-3.8. 

3.4-1.0 


3.8-4.1. 
4.0-4.3. 


length. 
Size - 


Smaller; largest specimen 107 mm. 
in standard length, usually 
much less than 100 mm. long. 


Larger; largest specimen 130 mm. 




long and smallest 107 mm. 



The posterior position of the pelvic fins and the few gill rakers 
readily separate patns from EyporhampTius unifasciatus (Ranzani), 
H. roherti (Valenciennes) {=hildebrandi Jordan and Evermann), 
H. snyderi Meek and Hildebrand, and JI. gilli Meek and Hildebrand, 
the other species reported from Middle America (Meek and Hilde- 
brand, 1923, pp. 236-241, pis. 16-17) . 

Acknowledgments. — Material of Eyporham/phus rosae was bor- 
rowed from the Stanford Natural History Museum, through the kind- 
ness of Miss Margaret Storey, and from the Chicago Natural History 
Museum, through the courtesy of K. P. Schmidt aiid Mrs. Marion 
Grey. I am grateful for this cooperation. 



LITERATURE CITED 

C'l'vu B. Gkoiii.k8 L. t". F. D. 

1.S17. Li' ri'pne aiiiiniil distribue d'apros son «irg;misiiti<Mi, \ynur scivir dc base 
ft lanatitmio coniparie, vol. 2. xviii + 532 pp., lllus. Pails. 
K\i:i{MAN.N. Bahton W.\rbkn, and Jknkins, Ouvkr Pkebi.es. 

ISStl. HeiMirt upon a (•«>llei-tion of flsh«'.s made at Guayiuas, Sonora, Mexico, 
with di'scriptioiis of new species. Pnic. U. S. Nat. Mas., vol. 14, pp. 
121-lt;.'^. 2 pis. 
GiM, Tii>:oi)oi;k Nicholas. 

\H'>9. Deseription of HiipoihitiiHihux. a new ^'enns of fislnss allied to Ilfttii- 

rhdniphu-s Cuv. Proc. Acad. Nat. Sci. Phil:idelphia, lSr>[), p. 131. 
18Js{. Note on the genera of Ileinirhaniphiri.tc Pi(.c. Acad. N.il. Sd. Pliila- 
delphia, ]8«3, pp. 272-273. 
IIi-RKK. Albert W. C. T. 

I!t44. A review of (In- lialfbeaks or Heniiramphidae of the Philippines and 
adjacent waters. Stanford Univ. Pnltl.. univ. ser.. binl. .sci., v«>l. 9, 
No. 2. pp. .S!^-8»5, 1 map. 
Hrmis. CaR[. Lkavitt, and Millkr. RoitKitr Rush. 

IIMI. DoroKOiita smUhi, the fiisl kmnvn gizzard shad from the Pacific drain- 
age of Middle America. Copeia, 11)41, No. 4. pi). 232-238, 1 fig. 
JoKDAX. Damd Stark, and GiiJirarr, Charles Henky. 

188<J. Description of a new species ot ncmirhamphwi (Hcmirhamphus rosae), 
from the coast of California. Proc. U. S. N.it. Mus., vol. 3, pp. 
33.v:«6. 
Meek, Seiu Eugknk. and G<)ss. Damd Kore. 

IXSii. A revi«-w of the American .spwies of (he genus n< niii lnunplitis. Proc. 
Acad. Nat. Sci. IMiiladelphia, 1S84. pp. 221-220. 
Mekk, Seth Eugene, and IIili)ebr.\m), Samuel Fheueruk. 

ir>23. The irfurine fishes of Panama : Part 1. Publ. Field Mus. Nat. Hist. No. 
21.";. zo.il. .ser.. vf.l. 1.'. xi + '■^^'^^ PP . 24 pis. 

PoKV. FkI.II'E. 

ista). Meniorias soImc I.i liistoria do i.i isi:i i\t ('\ih:i. vol. 2. f.-isc. 'J., pp. '.t7-33»>. 
S.Miin. .1. L. B. 

P»:'..''.. Tin' South African sixcies of the genus II< mil luiuiiiliiis <'uv. Trans. 
Roy. S(M-. South Africa, vol. 21, pt. 2. pp. 12!» l.'.d. 1 fig., 3 pis. 
Wk>.i>. Aii-rkd <'i.evela.m>. 

VXVA. Notes on fishes of the f.-millv I lemirhaiiii>hid.ic. Piihl. I'ifid Mas. Nat. 
Ilisl., zr.ol. s«'r.. vol. •_'2. itii. II <■><;. 

103 



u t •ovtmiiidiT ruiNTiNo orrict r it«i 



U S NATIONAL MUSEUM 



PROCEEDINGS VOL 9C PLATE 11 



u C_ 



y. 



y. 



;-> 



PROCEEDINGS 0.^ THE UNITED STATES NATIONAL MUSEUM 




issued I^^vA-sjIJ^b h '^« 



SMITHSONIAN INSTITUTION 
U. S. NATIONAL MUSEUM 



Vol. % Waihington: 1946 No. 3196 

NOTES Oy. RECENTLY MOUNTED REPTILE FOSSIL SKELE- 
TONS 'IN THE UNITED STATES NATIONAL MUSEUM 



By Charles W. Gilmore* 



Skeleton of a nearly complete Eocene crocodile and a partial skele- 
ton of Corythosaurus , a crested dinosaur of the Upper Cretaceous, have 
recently been added to the exhibition collection of fossil vertebrates in 
the United States National Museum. In the brief notes presented 
here, attention is called to some of the more interesting anatomical 
features of these specimens, and measurements arc given of all the 
more important bones in order to make these data available to stu- 
dents of the fossil Reptilia. The crocodile specimen appears to be 
unique in being the first complete skeleton to be mounted for exhibi- 
tion in this country. 

SKELETON OF CROCODILUS CLAVIS COPE 

Platks 12-1 f) 

Among the specimens collected by the 1930 Smithsonian Paleon- 
tologieal Expedition to the Bridger Basin in southwestern Wyoming 
was an unusually complete skeleton of a crocodile, U.S.N.M. No. 
127l'.». It was found by George B. Pearce in the badlands between 
lievett and Little Dry Creeks in horizon B of the liridger fornuition. 
The skeleton as it lay in the ground was only i)artially articulated, but 
inasmuch as it is an isolated specimen the few bones found detached 
and sf-attered can surely l)e regarded as peitaining to a single individ- 
ual. The skeletal parts preserved are as follows: Skull, lower jaws, 
23 presacral vertebrae, 2 sacral vertebrae, 33 caudal vcrtcbiac 17 
chevrons, complete pectoral and pelvic girdles, If) eer\ ical ribs. 17 

•Mr. Ollmorp died on ."^cptcmbor 27, 194'>.— Ki<. 

6W7.14— 16 1 Jtr) 



196 PROCEEDINGS OF THE NATIONAL MUSEUM ^^^ gg 

thoracic ribs whole or in part, right humerus, both radii, right ulnare, 
both radiaha, parts of 2 metacarpals, both femoro , both tibiae, both 
fibulae, both tarsi, all metatarsals, 1 1 phalangials, 2 inguals, and num- 
erous dermal scutes. 

On account of the rarity of good crocodilian skeL'tons in paleonto- 
logical collections, it seemed desirable to articulate this specimen for 
public exhibition. After several months' work this w;as accomplished 
by Norman H. Boss, chief preparator, who is to be higtdy commended 
on the excellent results achieved. 

The individual bones are thoroughly mineralized an d practically 
free from postmortem distortion. A few elements, ho'>vever, either 
through injuries or disease are abnormally deformed. The right 
scapula is a most interesting example of a badly healed fra cture. In 
life the scapula was cleanly broken through the narrowesu" part of 
the blade. This upper portion dropped down on the inside of the 
proximal half for fully an mch below the point of fracture, and inhere 
the two parts were securely knitted together by extraneous bvOny 
growth. Although this fracture must have been exceedingly painijil 
at the time, after healing the limb undoubtedly continued to function. 

The second metatarsal of the left hind foot exhibits a pathologic 
condition that has enlarged the shaft of the bone to nearly twice its 
normal size. This lesion can probably be attributed to an injury. 
Other lesions are found on the left coracoid, anterior thoracic ribs, 
caudal vertebrae, chevrons, and skull. That this animal was a pug- 
nacious individual and often engaged in combat, probably with 
others of its kind, is clearly indicated by the considerable number 
of healed wounds. 

There are 23 presacral vertebrae preserved, but a restored lumbar 
was introduced between the first and second, the only point showing 
evidence of a break in the series, in order to make the presacrals corre- 
spond in total number to the vertebral formula of Crocodilus americanus 
as determined by Mook.^ This introduction makes four lumbar 
vertebrae, whereas Mook recognizes only three in C. americanus; 
but as a mounted skeleton of this species (U.S.N.M. No. 14874) has 
four, it seems reasonable to assume that a similar variation may 
occur in the fossil species. 

The caudal series consists of 37 vertebrae of which 33 are original 
bones. Four vertebrae at the tip of the tail are fully restored. 
According to authorities the total number of caudal vertebrae is 
subject to considerable variation among living individuals of the same 
species. 

The skeleton, over all from tip to tip, measured a^long the curves 
of the spmal column has a length of about 9 feet 10 inches. 

Pending the publication of the monographic study of the Croco- 

1 Mook, C. C, Bull. Amor. RTus. Nat. Ffist., vol. 44, pp. 70-78, 1921. 

« 



REPTILE FOSSIL SKELETONS — GILMORB 197 

(lilia by Dr. C. C. Mook, this specimen is provisionally identified as 
portaiuin*]: to the species Croc(x!ih!.<t claris Cope. ITay * recoo7iizes 10 
species of (\ocodibt'i from the Bridger formation alone, but a thorough 
revision would doubtless greatly reduce this list. 

The mounted skeleton has been given a defiant attitude (see pi. 13) 
with the jaws agape in order better to display the mouthful of teeth. 
The pose and style of mount adopted were largely determined by the 
character of the skeleton. Because of the extreme hardness and 
brittleness of the fossilized bone, it was found impractical to drill 
the bones for secm'hig them to metal supports, and so Mr. Boss 
worked out a scheme of half relief and half free mount that overcame 
this difficulty an<l gives pleasing results, as is clearly shown in plates 
12 and lo. 

The skidl and lower jaws are unusualty complete and only slightly 
distorted by crushing. 

In the upper and lower mandibles there are alveoli for 76 teeth, 
premaxillaries 10, maxillaries 30, and dentaries 36. Of this dental 
series 29 teeth were found in place; enumerated from the front these 
were distributed as follows: Second of the right premaxillary, a germ 
tooth, and fourth of the left premaxillary; hi the right maxillary the 
fifth, nintli, tenth, and eleventh, none in the left maxillary; in the 
left dentaiy the first, fourth, fifth, sixth, seventh, (germ tooth), eighth 
(gemi tooth), ninth (germ tooth), tenth, and eleventh; in the right 
ramus the first, second, thii'd, fourth, fifth, sixth, seventh, eighth 
(germ tooth), twelfth, thirteenth, fifteenth, sixteenth, seventeenth, 
and eighteenth (the last four being germ teeth). 

The other teeth, some 39 in all, were found loose in the matrix 
surrounding the skull, and these have been arldtrarily inserted in the 
jaws. Thus of the 76 teeth forming the complete dental series, 
original tooth crowns of 69 are preserved. 

In Crocodilus, according to Mook,^ all the living species of the genus 
have 17 to 19 toclh in the upper sei-ios and oidy 15 in the lower, or a 
maximum total of 68 teeth in the mouth as contrasted with 76 in the 
extinct species. Of 4 other skulls fr<jiii the Bridger in the National 
Museum's collections none shows less than 36 in the lower dental 
series, and it is evident that the greater number of teelh constitutes an 
important feature for distingui.shing the extinct Kocene forms fr(»ni 
\]\(' extant member.^ of the genus. It also raises the question of the 
propriety of referring these Eocene crocodiles to the genus (Vorodllus. 
That , however, is outside the scope of the present paper, and no doubt 
will be fully considered by Mook in (he ronrse of his monogrHi)lii(; 
study of the order. 



> l\ny, f). I'., UiMlogrftphy onil catolfiRiio of ffissll vcrU'l)rnt<'s. Cnni'-rli' Itist. \\'ir liliirti.n V\ih]. sr*), 
pp. .■il2-.M3, 1928. 
• .Vlwjk. C. C. Hull. Amcr. Mus. Nat. ITIsl., vol. ii. p. I.M. 1021. 



198 



PROCEEDINGS OF THE NATIONAL MUSEUM 



The excellent preservation of the present skeleton offers an unusual 
opportunity for recording the important measurements of a single 
individual of an Eocene crocodile. In the tables that follow the prin- 
cipal measurements of the skeletal parts are given, following Mook's 



Table 1. — Comparative measurements {in mm.) of skull, pelvic, and limb bones in 

two species of Crocodilus 



Measurements 



Length of skull, tip of snout to supraoccipital 

Length of skull, tip of snout to ends of quadrates 

Breadth of skull, cranial table 

Breadth of skull, across fifth maxillary tooth 

Length of mandible 

Lp,ngth of scapula, total -. 

A nteroposterior diameter of superior border 

Anteroposterior diameter of inferior border 

Maximum thickness of distal end 

Length of coracoid, total --- 

Anteroposterior diameter, superior surface 

Anteroposterior diameter, inferior surface 

Length of humerus, total ..- 

Breadth of proximal end 

Breadth of distal end 

Circumference of shaft - 

Index of circumference over length 

Length of radius, totaL -.. 

Maximum diameter, proximal end 

Maximum diameter, distal end 

Length of ilium, total oblique - --- -- 

Length of ilium, total anteroposterior 

Distance across both ischiadic processes 

Maximum length of ischium, oblique 

Anteroposterior diameter, proximal end .- 

Maximum diameter, distal end 

Length of pubis, total _-_ _. 

Maximum diameter, proximal end 

Minimum diameter, proximal end 

Breadth distal end 

Length of femur, total 

Breadth , proximal end 

Breadth, distal end _ 

Circumference of shaft 

Distance from center of fourth trochanter to proximal end 
Distance from center of fourth trochanter to distal end 

T J i- (Center of fourth trochanter to proximal end} 

Index ratio; L- 

l Center of fourth trochanter to distal end J 

Length of tibia, total 

Maximum diameter, proximal end- __ 

Maximum diameter, distal end_ 

Length of fibula, total _ - 

Maximum diameter, proximal end 

Maximum diameter, distal end 

Circumference of shaf t_ _._ 

- , .. I Circumference of shaft] 

Index ratio; >._ 

I Total length 1 



C. clavis 
(U.S.N.M. 
No. 12719) 



418 
481 
110 
126 
535 
136 

77.3 

75.4 

27 
101 

60.4 

68.5 
180 

54 

50.5 

61 
338 
107 

26 

26.1 
125.6 
105 
145 
1.39 

49 

68 

93.5 

27 

17.4 

73.5 
223 

51.5 

51.5 

76 

81 
142 

0.570 

159 

43.1 

38 
156 

25 

24 

35 



C. americanus 

(A.M.N.H. 

No. 7139) 



1 Possibly an error, in view of greater proportions of most of the other measurements. 



REPTILE rOSSIL SKELETONS — GILMORE 



199 



system * used in doscribing: the osteology of the extant Orocodilvs 
aniericanvs. 

For convenience in reference I have included in parallel columns the 
measurements by Mook of a considerably larger (\ amcricanvs. 
In view of the antiquity of the extinct skeleton these measurements 
show a remarkable similarit)" of proportions between the living and 
extinct forms. 

Tahle 2. — Comparative measurements {in mm.) of cervical, dorsal, luvihar, and 

sacral vertebrae 





Length of 
centrum ' 


Breadth of 

centrum, 

anterior end 


Spread of 

preiygapo- 

pliyses 


Spread of 
postzyga- 
pophyses 


Spread of 

diapophy- 

ses 


Total 
height 


Vertebrae 


o 

s 

6 

•z 

S 

03 


d 

< 


g 
d 

s 
z 

CO 

t> 


1 

K 
Z 


OJ 

B 

d 
Z 

z 

DO 


2 
d 

z 

w 
z 

< 


d 

z 
z 

00 


d 

z 

a 
z 


§ 

d 

z 

i 
z 

vi 


o 
d 

z 

a 
z 


d 

z 
z 

CO 


d 
Z 

w 
z 


Cervical 2 


4g.6 

34 

35 

32.5 

31.9 

32.5 

31 

32 

32 

34.9 

36.0 

37 

39 

39 

39 

41 

41 

41 

41 

39.9 


04.5 

67 

68 

66 

65 

64 

65.6 

63 

65 

66 

08 

68 

71 

73 

73 

74 

76 

78 

79 

78 

77 

77 

09 

59 

62 


25 

25.5 

27 

31 

30.7 

34 

32 

33 

33 

33 

33.9 

32.8 

34 

35 

35 

30 

36.2 
33.6 
44 
30 


46 
40 
43 
44 
44 
52 
60 
54 
50 
53 
50 
48 
49 
49 
60 
60 
61 
57 
55 
65 
60 
56 
65 
60 
60 


28 

32.2 

41 

41 

50 

48.5 

65 

54 

63 

49 

00.5 

GO. 6 

60 

61 

68 

61 

62 

63 

55 

44 

30 


30 

40 

44 

54 

61 

64 

68.5 

68 

66 

07 

76 

82 

84 

82 

80 

81.5 

80 

80 

SO 

83 

82.6 

86 

88 

87 

65 


26 
41 

48 

40 

52.5 

55 

62.5 

58 

00 

59.5 

58,5 

50 

62 

54 

59.5 

55.4 

54 

33.8 
28 
42.6 


33 

41 

49 

54 

58 

64 

63.5 

62 

63 

70 

79 

81 

79 

76 

78 

78 

70 

83 

81 

81 

81 

85 

84 

62.6 

53.5 


51.7 

41 

43 

51 

49 

00 

73.3 

84 
101 
105 
127 
149 
159 
150 
160 
100 
100 
UK) 
144 
130 


53 

35.5 

49 

55 

58 

63 

64.5 

03 

61 

CO 
107 
150 
183 
200 
213 
218 
230 
242 
276 
277 
250 


79 
88 
06 

105 

122.9 
121.6 

118 
95 
90 
88.5 
85 
80 
81 
79 
80 

83 

88 
79.8 


87 


3 


127 


4 


140 


6 


156 


6 


166 


7 


167 


8 


175 


Dorsal 1 ' 


180 


2 


185 


3 


180 


4 


181 


5 


100 





150 


7 


148 


8 


145 





140 


10 


140 


11 

12 


136 
139 


r.iimbar 1 


134 


2 


133 


3 


39 
38 
42 
43.7 


120 

156 
110 


235 
218 
230 
202 


132 


i 

Sacral 1 


132 
135 


2 


142 







' Th*> mntKurriiicnts of the vertebrnl centra of Ihn presacral st-rlr* of the fo.ssil specimen do not Include 
the ball, and so In using these mea.surenicnts allowance must bo made for this omission. This discrepancy 
was dui) to the fact that this prjrtlnn of the backbone was articulated in nxed position before It was decided 
to tiko thi? scries of mcfi.<iuremonts. 



•Ibid., pp. 71-100. 



200 



PROCEEDINGS OF THE NATIONAL MUSEUM 



Table 3. — Comparative measurements {in mm.) of caudal vertebrae 






Length of 


Breadth of 


Height of 


Spread of 


Spread of 


Height 


Spread of 




centrum, an- 


centrum, an- 


prezygapo- 


postzygapo- 


transverse 








terior end 


terior end 


physes 


physes 






processes 




o» 


OS 


05 


Ol 


OS 


OS 


OS 


OS 


OS 


OS 


OS 


OS 


OS 




































































r-" 
























































Caudals 


6 


o 


d 


o 


6 


o 


d 
A 


o 
A 


d 
•A 


o 


d 
'A 


1 


d 
"A 


o 
A, 




S 


w 


^ 


w 


s 


w 


i, 


te 


i 


w 


i 


w 


% 


te 




^^, 


Z 


Z 


A 


iu 


2 


"A 


A 


'A 


'A 


Z 


'A 


"A 


A 




ai 


% 


m 


:§■ 


m 


s 


CO 


% 


m 


S 


m 


<? 


CO 


s 




P 


< 


tD 


<i 


t2 


< 


t) 


< 


t> 


< 


P 


■< 


ID 


< 


1 


55.9 


73 


34.3 


51 


29 


46 


45 


58.5 


34 


52 


83.9 


130 


122 


196 


•2 


51.7 


69 


32 


50 


30 


43 




59.5 


30 


48 


77 


134 


132 


195 


3 


51 


70 


29.4 


45.5 


31.5 


44 




54 


27 


43 


83 


135 


126 


186 


4 


52 


73 


27 


46 


31 


42 




50 




40 


79 


135 


116 


175 


5 


55 


75 


27.4 


44.6 


28.6 


41.5 


27.8 


46.5 


26 


38.5 




136 




166 


6 


55.7 


76 


26.9 


43 


27 


38.5 


28 


45 


21 


36 


75.5 


130 


112 


157 


7 


53.7 


67 


25 


40.5 


24 


37 


26 


42 


21 


32 




132.5 




141 


8 


53.2 


77 


25 


44 


25 


44 


25 


37 


22 


29.5 




118 




130.5 


9 


53.5 


68 


23 


34.5 


23 


36.6 


28 


37 


20.9 


32 


67.5 


103 




120 


10 


53 


78 


24.4 


35.5 


24.1 


33 


24 


36.5 


19 


32 


68.8 


101.5 




118 


U 


53 


77 


23 


33.5 


23 


30.5 


27 


38 


21.5 


27 


71 


100 




120 


12 


54.3 


74 


22 


32 


22.5 


29 


22 


34 




27 


67 


107 




107 


13 


54 


78 


22.5 


32 


22 


28 




33 




23.5 




104 




88.5 


14 


53.4 


75 


21.7 


31 


23 


26 




28 




25 


66 


102 




44.5 


15 


52 

52 

50 

51 

50 

49 

49.8 

47.5 

46 

44.8 

43.6 

42.5 

40.5 

39 

30.5 


72 

75 

74.5 

73 

70.5 

70 

66 

66 

65 

63 

60.5 

59 

57 

53 

49 

46 

46 

34 

34 


21.5 

19.5 

19 

20 

18 

17 

18 

16 

15 

14.5 

13.5 

12 

7 

6.3 

9 


29.5 

28.5 

27 

25 

20.4 

22.5 

21 

20 

18 

20 

16 

15 

13.5 

13 

12 

10.5 

10 

9 

7.5 


20.5 

21 
21 
18 
18 

16.6 
16 
16 

15.2 
15 
14 

13.1 
7.5 

7 


26 
26 
24 
25 
23 
22 
20 
20 
18 
18 
17 
15 
14 
12 
11 
10 
9 
8 
11 


15 
14 

14 

12.5 
11.2 
10.5 

7 

6.9 

6 


28 

28 

25 

22 

19 

18 

15 

14.5 

12.5 

10.5 
9.5 
8 

6.5 
5.5 
3.5 
2.5 
3 

•4.5 


10 
7 

7 

6 

4 

4.5 

4 

4 

3:9 

3 

3 


24.5 

21.5 

18 

18 

15 

11 

11.5 

7.6 

6 

3.5 

3.6 

2.5 

1 

1 

1 
.5 

3 
12.5 

1 


69 

65.5 
65 

46 
45 
38.9 


102 
106 
105 
118 
109 
88 
96 
87 
82 
65 
50 
50 
42 
35 






16 






17 






18 






19 






20 






21 






22 






23 






24 






25 






26 






27 






28 






29 






30 


27 
22.5 






31 






32 






33 




13.5 



















1 Estimates 



NOTE ON DISEASED CROCODILE VERTEBRAE 



Plate 16 

A second crocodile specimen, U.S.N.M. No. 12990, from the 
Bridger, Eocene, consisting of the greater portion of a skeleton, 
shows a pathologic condition of two dorsal vertebrae that is almost 
identical with the lesion described by Moodie ^ on the caudal verte- 
brae of a sauropodoiis dinosaur from the Jurassic. 

It is a spongy growth that surrounds the intervertebral articular 
surfaces extending well outward on the sides of both centra (see 

5 Moodie, Roy L., Amer. Journ. Sei., vol. 41, pp. 530-531. 19IP. 



REPTILE FOSSIL SKELETONS — GILMORK 201 

pi. IG). It entirely encircles the centra and has involved two- 
thirds of the two bones. All evidence of separate structure is prac- 
tically obliterated. The growth is cjuite symmetrical on the two 
sides. 

Moodie in the case of the lesion on tiie dinosaur caudals says: 
"The enlargement is somewhat suggestive of the lesion of chronic 
osteomyelitis. It may be a callous growth due possibly to a fracture 
of the caudal vertebrae; or it may be a bone tumor." Its true 
nature is, of course, uncertain, but mention is made of this specimen 
here in order to call it to the attention of students of modern pa- 
thology who may be interested in the study of the nature and origin 
of disease. 

RufTer' has reported typical lesions indicating spondylitis dejor- 
mans in the vertebrae of the Miocene crocodile Tomistoma dowsoni, 
from Egypt. In this specunen the extraneous osseous tissue, obvi- 
ously pathologic, binds the vertebrae together. The new bone, 
however, is thicker on one side than on the other, and Moodie ^ 
observes that "in the crocodile as in man the disease is more marked 
on one side." The symmetrical nature of the lesion in the National 
Museum crocodile shown in plate 16 would therefore rule out spon- 
dylitis dejormans as being responsible for the development of this 
altuormality. 

ON A SPECIMEN OF CORYTHOSAURUS 

Plates 17-19 

A partial skeleton of Corythosaurus recently added to the exhibition 
scries in the United States National Museum consists of the complete 
articulated tail, pelvis, hind limbs, and feet, with several small patches 
of skin impressions and o-ssified tendons. This specimen is mounted 
in rehef so as to display the right side; the sandstone blocks containing 
the bones have been assembled in the same relationships the}' occu- 
pied in the ground. The preparation ant! mounting were done by 
Norman H. Boss, and it was due to liis skillful manipulation that so 
much of the epidermal impressions were preserved. 

The specimen (L'.S.X.M. No. 1549:^) was acquired by purchase 
from the Royal Paleontological Museum of the University of Toronto 
for use in connection with the Smithsonian exhibit at the Texas 
Centennial Exposition at Dallas in \\)'M . It was collected l)y Levi 
Sternberg from the Belly River formation, rpjjer Cretaceous, 2 miles 
south of St(!veville, on the Red Deer River, Alberta, Canada, in VX.V.\. 

From an exhibition standpoint this si)eeimen is of interest in having 
several patches of skin impressions preserved. When found the 

• HufT( r, i^\t Marc Arnian<l, A pftlhoIoRlcnl sih-cIiiu'ii diillriK from tho Lower Mio«'iio imtIik!. ]n Appon- 
(llx to Foiirtati's "C"onlrl»>iillon k \'t,i\ii\n Vfrt6br*s .Mloc^kmit do rfegypto," pp. 101-109, illiis. Survey 
Dopartnient, MInhtry of FInanro, ralro, 1030. 

' Moo'llo, U. L., ralropatliolo?) , p. 17.^, 1OT3. 



202 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

skeleton was lying on its left side, but the definition of the skin pat- 
tern of that side had been dulled or wholly destroyed by the presence 
of a considerable amount of vegetal matter on which it lay. The 
bedding plane beneath the skeleton was unusually in-egular, indicat- 
ing that the cross-bedded planes were laid down by currents acting 
from different directions. 

The caudal series consists of 75 vertebrae and appears to be com- 
plete. This information, together with that furnished by Brown,^ 
shows the complete vertebral formula of Corythosaurus to be 15 cer- 
vicals, 19 dorsals, 8 sacrals, and 75 caudals. It is presumed, how- 
ever, that as in many other reptiles the caudal series will be subject 
to some individual variation in number. The first 16 caudal verte- 
brae have transverse processes as in the type. The first chevron on 
the tail of Corythosaurus is carried between the fifth and sixth caudal 
vertebrae; thus the total number of caudal vertebrae can be accu- 
rately determined and a close estimate of their combined length can 
be obtained from this specimen which has the first two caudals com- 
pletely hidden by the overlying ihum and a patch of skin impressions. 
Measured along the curve this tail has a complete length of about 455 
cm. (14 feet 11 inches). 

A complete description of the ossified tendons of Corythosaurus has 
already been given by Brown, and thus it is only necessary to men- 
tion that the present specimen fully corroborates his determination 
that they are disposed in two layers. 

Small patches of skin impressions are present on the midsection of 
the tail, on the pelvis, and on the feet. Those pieces of the integu- 
ment best preserved cover the thirty-first to the thirty-fourth caudal 
vertebrae, respectively. The detailed mosaic pattern of the flat, 
polygonal scales is clearly and beautifully shown in plate 18. Origi- 
nally the whole midsection of the tail beginning with the sixteenth 
caudal was covered by skin impressions, but most of the center of this 
patch was so friable that it could not be preserved. The outside 
portions, however, outline the original width of the tail at this 
point. The skin on and below the right ilium is composed of scales, 
slightly larger than those of the midcaudal region, but otherwise 
they seem to be indistinguishable. The pattern of the scales on 
the feet is dim and illy defined and adds nothing to oiu- previous 
knowledge. 

In the course of preparing this specimen many small detached pieces 
of skin were found in the matrix. Several of these were folded and 
others had been completely reversed. Six species of this genus have 
been named, aU from the Belly River formation of midwestern Canada. 



I 8 Brown, Barnum, Bull. Amer. Mus. Nat. Hist., vol. 35, p. 710, 1916. 



REPTILE FOSSIL SKELETONS — GILMORE 



203 



Listed in chronological order these are: C. casuarius Brown,' C. 
excamtus Gilmore,"^ C. infermeduis Parks," C. bicrisfatus Parks,'^ 
0. Jronfalis Parks,'* and C. breincristatus Parks.'* 

Five of the six species were established on skull characters alono 
and are distinguished chiefly by differences found in the shape and 
extent of the crest. In the absence of the skull in the specimen under 
consideration it appears quite impossible at this time to make a definite 
identification of the species. However, on the basis of similarity of 
skin pattern and close agreement in proportions to the type specimen 
of Con/thosaurus casuarius, as shown in table 4, this specimen is 
provisionally identified as pertaining to that species. 

Table 4. — Comparative measuremenls (in cm.) of (u'o specimens of Corythosaurus 

casuarius 



Measurements 



U.S.N.M. 
No. 15493 



A.M.N.n. 
No. 5338, typo 



Length ot longest che\Ton 

Ischhim, greatest length 

Ischium, length of terminal foot 

Femur, greatest length 

Femur, position of fourth trochanter from central point to top of femur 

Tibia, length of tibia and astragalus 

Fibula, length 



40 
113.6 

20 
115.9 

58.5 
100.7 

98.2 



38.5 
lfr3 

22 
lOS 

58 
100 

95 



» Brown, Bamum, Bull. Amer. Mus. Nat. Hist., vol. 33, pp. 559-565, pi. 41, 1914. 
'• Ollmore, C. W., Can. Field-Nat., vol. 37, pp. 46-52, 1923. 

" Parks, W. A., Univ. Toronto Stud., geol. scr.. No. 15, pp. 1-57, 13 Ogs., pis. 1-4, 1923. 
n Parks. W. A., Univ. Toronto Stud., gool. scr.. No. 37, pp. 2^45, pis. 4-8, 1935. 



u t. aovcmiMCNT miNTiNa orricii it4« 



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U. S. NATIONAL MUSEUM 



PROCEEDINGS, VOL. 96 PLATE 13 







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PROCEEDINGS. VOL. 96 PLATE 14 




U. S. NATIONAL MUSEUM 



PROCEEDINGS, VOL. 96 PLATE 15 




2: 



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U S. NATIONAL MUSEUM 



PROCEEDINGS. VOL. 96 PLATE 16 










Diseased dorsal vertebrae of Crocodilus sp. (U.S.N. M. No. 12W0): Ipper liK"re, \eiilial 
view; lower lig»rp, lateral view. liotli natural size. 



U. S. NATIONAL MUSEUM 



PROCEEDINGS, VOL. 96 PLATE 17 




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U S NATIONAL MUSEUM 



PROCEEDINGS, VOL 96 PLATE 18 




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PROCEEDINGS. VOU. 96 PLATE 19 




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PROCEEDINGS OF TME UNITED STATES NATIONAL MUSEUM 



iistiej ''- ''' - ' "a^l hy the 




SMITHSONIAN INSTITUTION 

U. S. NATIONAL MUSEUM 



Vol. 96 Wa.hington: 1946 No. 3197 

THE ONYCHOPHORES OF PANAMA AND THE 
CANAL ZONE 



By Austin H. Clark and James Zetek 



For the past dozen years the junior author has been interested in 
colli'ctin'4 and obsorvinj]: the several species of Onychophora occurring 
in Panama and the Canal Zone. He sent to Hie senior author a 
collection including 82 specimens from various localities, among which 
were representatives of all but one (Peripatus ruber) of the species 
recorded from the region; this omission, however, was more than 
compensated by the presence of two species {Oroperipatus eisenii and 
Epiperipatus biolhyi) not previously known from this area, although 
the first has been rej^orLed both from farther north and from farther 
south. 

The number of species of Ouyc^hophora now known from Panama 
and the Canal Zone is unusually large for a region of such limited 
extent, amounting to no less than seven, distributed in four genera. 
These seven species are: Oroprripntiis eisfuii, 0. corradi, Macrnperi- 
patus gcayi, Peripatus ruber, Eplpcripaius brasilicnsis, E. edwardsii, and 
E. biolleyi. 

The richest locality was at El Cermeuo, where four species, Ornpcri- 
pntufi ei.Hfnii, 0. corradi, Iipiperijuilus cdvdrdsii, and E. bifdleyi, were 
found. Here the coconut habitat was the most productive, and they 
were ratlier (•onimon. These j)ulms were not more than 5 years old, 
and the sheaths of the fronds wer(! usually a foot or two from tiio 
ground. All it was necessary to do was to pull off these dead sheaths. 

M8730— 40 205 



206 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

It is quite damp where the onychophores are found. There were not 
many rocks or boards on the ground, which may account for the 
presence of these creatures under the coconut-palm sheaths. 

From Barro Colorado Island three species are now known, Oroperi- 
patus corradi, Macroperipatus geayi, and Epiperipatus brasiliensis var. 
vagans. 

Dr. Waldo L. Schmitt has found onychophores in the dry season 
under rocks where the soil remained moist in a dry stream bed, and 
0. eisenii and E. biolleyi have been found under stones by the sides of 
streams. One species has been found in bromelias in Central America, 
though not in Panama. 

As the records of the species of Onychophora known from Panama 
and the Canal Zone are widely scattered, it has seemed worth while to 
include all of them herein, together with a key for their identification 
and a bibliography. 

KEY TO THE ONYCHOPHORA OCCURRING IN PANAMA AND THE 

CANAL ZONE 

a*. Urinary papilla of legs IV and V included in third arc of ambulatory pad ; 

4 foot papillae, 2 on each side of foot. 

51. Urinary papillae of legs IV and V dividing third arc into 2 segments of 

which the posterior is smaller than the anterior, though much broader 

than long; urinary tubercle wholly united to anterior portion of arc; 

26-29 (usually 28) pairs of legs in females, 24-25 (usually 25) in males; 

length 14-39 mm Oroperipatus corradi 

b^. Urinary papillae of legs IV and V dividing third arc into 2 segments of 
which the posterior is very small, scarcely broader than long; urinary 
tubercle more or less independent of anterior segment; 27-29 (usually 
28) pairs of legs in females, 23-26 (usually 25) in males; length 13-70 mm. 

Oroperipatus eisenii 

a^. Urinary papilla on legs IV and V below third arc, deeply indenting fourth; 

3 foot papillae, 2 anterior and 1 posterior. 

¥. Papillae of dorsal surface each on an oblong or squarish base, the oblong 

bases elongated in direction of long axis of animal; 30-33 pairs of legs 

in females, 28 in males; length 27-100 mm Macroperipatus geayi 

¥. Papillae of dorsal surface on highly irregular bases, the plications usually 
appearing undivided. 
c^ Principal papillae of dorsal surface of very different sizes, some very 
conspicuous and cylindrical, others smaller and conical, usually 3 of 
the smaller between 2 of the larger; papillae separated by rather broad 
intervals in which accessory papillae occur; males usually with crural 
tubercles on more than 2 pregenital pairs of legs; 29-30 (usually 30) 

pairs of legs in females; length 29-52 mm Peripatus ruber 

c*. Principal papillae of dorsal surface all of same type, passing through all 
intermediate stages from large to small; papillae closely set, though 
with occasional accessory papillae between them; in small individuals 
some of the papillae predominant; crural tubercles of males on 2 
pregenital pairs of legs. 
dK On dorsal surface above each pair of legs a few short incomplete plica- 
tions, tapering to a point on each side intercalated between the others. 



ONYCHOPHORES OF PANAMA CLARK AND ZETEK 207 

eK Fourth arc on logs IV and V strongly arclied beneath urinary papilla, 
but not divided into segments; urinary papilla attachal to third 
arc by a narrow band at deepest point in incision in latter; 29-34 
pairs of legs in females, 28-30 in males; length 23 5G mm. 

Epiperipatus edwardsii 
c*. Fourth arc on legs IV and V broken into 2 or 3 well-separated unequal 
parts; urinary papilla wholly independent of third arc; 30 pairs of 
legs iu females, 26-2S in males; length 25-3(5 mm. 

Epiperipatus bioUeyi 

d-. No short incomplete plications on the dorsal surface visible in dorsal 

view, the plications appearing wholly regular; 31-32 pairs of legs in 

females, 29 in males; length 37-80 mm. Epiperipatus brasiliensia 

Genus OROPERIPATUS Cockerell 

OHOPEHIPATUS EISENH (Wheeler) 

Peripatus eisenii Wheeler, Journ. Morph., vol. 15, pp. 1-8, pi. 1, 1898 (Tepicj 
Mexico). — FcimMANN, M^m. Soc. Neuchfit. Sci. Nat., vol. 5, pp. 176-192, 
1912 (Rio PurtSs, Brazil). 

Oroperipatus eiseni A. II. Clakk, Proc. Biol. Soc. Washington, vol. 26, p. 16, 1913 
(listed). 

N'ew records.— lEA Cermeno, Panama; J. Zctek, July 8, 1941. Fe" 
males: (1) 55 mm. long;, 4 mm. broad, 28 pairs of legs. (2) 52 mm. 
long, 3.5 mm. broad, 28 pairs of legs. (3) 47 mm. long, 3 mm. broad, 
28 pairs of legs. (4) 46 mm. long, 3 mm. broad, 27 pairs of legs. 
(5) 45 mm. long, 3 mm. broad, 29 pairs of legs. (6) 45 mm. long, 
2.5 nmi. broad, 28 pairs of legs. (7) 42 mm. long, 2.5 mm. broad, 
28 pairs of legs. (8) 37 mm. long, 2 mm. broad, 29 pairs of legs. 
(9) 30 mm. long, 2 mm. broad, 28 pairs of legs. 

El Cermeno, Panama; J. Zetek, July 15, 1941. Females: (1) GO 
mm. long, 3.5 rajn, broad, 28 pairs of legs. (2) 59 mm. long, 3.5 mm. 
broad, 28 pairs of legs. (3) 55 mm. long, 3.5 mm. broad, 28 pairs of 
legs. (4) 53 mm. long, 3.5 mm. broad, 28 pairs of legs. (5) 52 mm. 
long, 3.5 mm. broad, 28 pairs of legs. (6) 50 mm. long, 3.5 mm. broad, 
28 pairs of legs. (7) 48 mm. long, 4 mm. broad, 28 pairs of legs. 
(8) 43 mm. long, 4 mm. broad, 28 pairs of legs. Mule: (1) 30 mm. 
long, 2 mm. broad, 25 pairs of legs. 

El Cermeno, Panama; J. Zetek, August 5, 1941. Females: (1) GO 
mm. long, 3.5 mm. broad, 28 pairs of legs. (2) 55 mm. long, 3 mm. 
broad, 28 pairs of legs. 

Panama City, Panaiiiii ; J. Zetek, Scplcnibcf 1939. Females: 
(1) 70 mm. long, 3.5 mm. broad, 28 pairs of legs. (2) ()9 mm. long, 
4 mm. broad, 28 pairs of legs. (3) Go mm. long, 3.5 mm. broad, 28 
pairs of legs. (4) 04 mm. long, 3 mm. broad, 28 pairs of legs. (5) 
61 mm. long, 3.5 mm. broad, 2S i)airs of legs. (G) (10 niiu. long, 3 mm. 
broad, 28 pairs of legs. 

Range. — Mexico (Tej)ie); Puiuiiua {VA Ceiiinno, rmuima City); 
Brazil (Kio Pin us). 

Note. — At El Cermeno this species was found in situations similar 
to those favored by 0. corradi. 



208 PROCEEDINGS OF THE NATIONAL MUSEUM vol.98 

OROPERIPATUS CORRADI (Camerano) 

Peripatus corradi Camerano, Boll. Mus. Zool. Anat. Comp. Univ. Torino, vol. 13, 
No. 316, pp. 2, 3, 1898 (Ecuador). — Bouvier, Ann. Sci. Nat., ser. 9, zool., 
vol. 2, p. 120, pi. 3, fig. 15, pi. 4, figs. 29, 30, text figs. 6, p. 15, 18, p. 20, 42, 
p. 38, 63, p. 124, and 64, 65, p. 125, 1905. 

Oroperipatus corradoi A. H. Clark, Proc. Biol. Soc. Washington, vol. 26, p. 18, 
1913 (listed); Smithsonian Misc. Coll., vol. 63, No. 2, p. 1, 1914 (Ancon, 
Canal Zone; notes); Zool. Anz., vol. 45, No. 4. p. 146, 1914 (Ancon). — Fuhr- 
MANN, Abh. Senck. naturf. Ges., vol. 36, Heft 2, pp. 277-283, 1915.— Brues, 
Psyche, vol. 32, No. 3, p. 159, 1925 (Canal Zone). 

New records. — El Cermeno, Panama; J. Zetek, July 8, 1941. Fe- 
males: (1) 32 mm. long, 2 mm. broad, 28 pairs of legs. (2) 30 mm. 
long, 2.5 mm. broad, 28 paii's of legs. (3) 28 mm. long, 2.5 mm. 
broad, 27 pairs of legs. (4) 25 mm. long, 1.5 mm. broad, 28 pairs of 
legs. (5) 16 mm. long, 1.5 mm. broad, 28 pairs of legs. (6) 15 mm. 
long, 1 mm. broad, 29 pairs of legs. Males: (1) 31 mm. long, 2 mm. 
broad, 25 pairs of legs. (2) 30 mm. long, 2 mm. broad, 25 pairs of legs. 
(3) 28 mm. long, 2 mm. broad, 25 pairs of legs. (4) 28 mm. long, 2 
mm. broad, 25 pairs of legs. (5) 27 mm. long, 2 mm. broad, 25 pairs 
of legs. (6) 27 mm. long, 2 mm. broad, 25 pairs of legs. (7) 26 mm. 
long, 1.5 mm. broad, 25 pan-s of legs. (8) 23 mm. long, 1.5 mm. 
broad, 25 pairs of legs. (9) 18 mm. long, 1.6 mm. broad, 25 pairs of 
legs. (10) 18 mm. long, 1.5 mm. broad, 25 pairs of legs. (11) 18 mm. 
long, 1.5 mm. broad, 25 pairs of legs. (12) 17 mm. long, 1.3 mm. 
broad, 25 pairs of legs. (13) 15 mm. long, 1 mm. broad, 25 paiis of 
legs. 

Barro Colorado Island, Canal Zone; J. Zetek, August 1933. Males: 
(1) 33 mm. long, 2.5 mm. broad, 25 pairs of legs. (2) 30 mm. long, 
2.5 mm. broad, 25 pairs of legs. 

Range. — Ecuador (Quito, Balzar, Guayaquil); Canal Zone (Ancon, 
Barro Colorado Island); Panama (El Cermeno). 

Notes. — At El Cermeno this species was found between the broad 
sheaths of old fronds and the trimks of coconut palms roughly 5 years 
old, and also under boards resting on the ground. On Barro Colorado 
Island it was usually met with under logs and stones and occasionally 
with the ground-termite stakes. At Ancon it was found in earth to 
the depth of about 1 foot about roots of papaya. 

Genus MACROPERIPATUS A. H. Clark 

MACROPERIPATUS GEAYI (Bonvier) 

Peripatus geayi Bouvier, Comptes Rendus Acad. Sci. Paris, vol. 128, p. 1345, 

1899 (French Guiana); Ann. Sci. Nat., ser. 9, zool., vol. 2, p. 200, pi. 6, 

figs. 42, 43, text figs. 1, p. 36, and 86, p. 203, 1905. 
Macroperipatus geayi A. H. Clark, Proc. Biol. Soc. Washington, vol. 26, p. 17, 

1913 (listed) ; Smithsonian Misc. Coll., vol. 63, No. 2, p. 2, 1914 (from Clark, 

1913); vol. 65, No. 1, p. 23, 1915 (from Clark, 1913). 



OXYCHOPHOHES OF PANAMA — CLARK AND ZETEK 209 

Peripalus (Mncroperipolus) geayi A. II. Clauk, Smilhsoiiian Misc. Coll., vol. GO, 
No. 17, pp. 1-5, 1913 (La Chorrera, Panama; notes). — BurKS, Psyche, vol. 
32, No. 3, p. 160, 1925 (from Clark, 1913). 

New records. — Barro Colorado Island, Canal Zone; J. Zrtok, July 
1941. Females: (1) 100 mm. long, 8 mm. broad, 33 pairs of legs. 
(2) 70 mm. long, 4.5 mm. broad, 31 pairs of legs. (3) 70 mm. long, 
4 mm. broad, 30 pairs of legs. (4) 64 mm. long, 4 mm. broad, 30 
pairs of legs. (5) 38 mm. long, 3 nmi. broad, 33 pairs of legs. 

Pedro Miguel, Canal Zone; J. Zetek, April 1938. Females: (1) 84 
mm. long. 5 mm. broad, 32 pairs of legs. (2) 59 mm. long, 3 mm. 
broad, 31 pairs of legs. Afalcs: (1) 30 mm. long, 2 mm. broad, 28 
pairs of legs. (2) 27 mm. long, 1.5 nmi. broad, 28 pairs of legs. 

Balboa, Canal Zone; J. Zetek, 1944. Female: (1) 70 mm. long v/itli 
33 pairs of legs. 

Range.— French Guiana; Colombia; Panama (La Chorrera); Canal 
Zone (Barro Colorado Island, Pedro Miguel, Balboa). 

Notes. — On Barro Colorado Island tliis species occurred under 
fallen logs and in leaf mold, at Pedro Miguel mider stones and boards 
on the ground. Large onychophores from the Orsini citrus orchard 
clo<e to La Campana were collected under fallen logs and under stones. 
These were not seen by the senior author but were presumably of this 
species. 

Genus PERIPATUS Guilding 

PERIPATUS RUBER Fuhrmann 

Peripatua ruber Fuhrmann, M6m. Soc. NeucMt. Sci. Nat., vol. 5, p. 190, 1912 
(Rancho IJcdondo, Costa Rica) ; Zool. Anz., vol. 42, No. 6, p. 247, figs. 12-14, 
p. 248, 1913 (redescribed). — A. II. Clark, Smithsonian Misc. Coll., vol. 05, 
No. 1, p. 24, 1915 (Lino; from Clark, 1914).— Fuhrmann, Abh. Senck. 
naturf. Ges., vol. 30, Heft 2, pp. 277-2^3, 1915. 

IPeripalus {Epipcripalua) biollciji var. brthcli Cockerkll, Proc. Biol. Soc. Wash- 
ington, vol. 26, p. 87, 1913 (Puerto liarrios, Guatemala). 

Peripalus (Peripatus) ruber A. II. Clark, Zool. Anz., vol. 45, No. 4, p. 145, 1914 
(Lino, near Bouquete, Province of Chiricjuf, Panama; 4,100-4,500 feet; notes 
on 3 specimens). 

Range. — Costa Kica (Kancho Kedondo); Panama (Lino, near Bou- 
quete, Province of Chiriquf, 4,100-4,500 feet); ?Guatemala (Puerto 
Barrios). 

N^ote. — From the desciiption l-*rofessor Cockerell's Pcriyatus {Kpi- 
peripatus) biolleyi var. betfiell appears to be Peripalus ruber. His typo 
specimen is not at present available for reexamination. 

Genus KPIPEKIPATUS A. H. Clark 

EPIPERIPATUS BRA.SILIEN»LS (Bouthir) 

Peripatua braailienaia Bouvier, Comptes RenduH Acad. Sci., Paris, vol. 129, p. 
1031, 1899; Ann. Sci. Nat., scr. 9, zool., vol. 2, p. 269, pi. 7, fig. 63, 



210 PROCEEDINGS OF THE NATIONAL MUSEUM vol.06 

pi. 8, figs. 64, 65, text figs. 100-103, p. 273, 1905 (Santar^m; ?San Pablo, 
Panama); Bull. Soc. Philomatique, ser. 9, vol. 10, pp. 50-52, 1908 (M^rida, 
Venezuela; San Pablo, Panama, confirmed). — Fuhrmann, M^m. Soc. 
Neuchat. Sci. Nat., vol. 5, p. 190, 1912 (Brazil and Guiana to Panama), 

Epiperipatus brasiliensis A. H. Clark, Proc. Biol. Soc. Washington, vol. 26, p. 18, 
1913 (listed); Smithsonian Misc. Coll., vol. 63, No. 2, p. 2, 1914 (listed from 
Panama); vol. 65, No. 1, p. 23, 1915 (San Pablo, Panama; from Bouvier). — 
Fuhrmann, Abh. Senck. naturf. Ges., vol. 36, Heft 2, pp. 277-283. 1915. 

Peripatus (Epiperipalus) brasiliensis var. vagans Bruep, Psj'che, vol. 32, No. 3, 
p. 162, 1925 (Barro Colorado Island, Las Cascadas, Fort Sherman, and 
Chinilla, Canal Zone; Rio Tapia, Panama; description). 

New records. — Barro Colorado Island, Canal Zone; J. Zetek, 
January-May, 1942. Females: (1) 50 mm. long, 3.5 mm. broad, 33 
pairs of legs. (2) 40 mm. long, 3 mm. broad, 33 pairs of legs. (3) 

30 mm. long, 2 mm. broad, 31 pairs of legs. (4) 23 mm. long, 2 mm. 
broad, 33 pairs of legs. Males: (1) 30 mm. long, 2.5 mm. broad, 29 
pairs of legs. (2) 18 mm. long, 2 mm. broad, 29 pairs of legs. (3) 
17 mm. long, 2 m^m. broad, 29 pairs of legs. 

Barro Colorado Island, Canal Zone; J. Zetek, August 1933. Male: 
42 mm. long, 2.5 mm. broad, 29 pairs of legs. 

Balboa, Canal Zone; J. Zetek, 1944. Females: (1) 55 mm. long, 32 
pairs of legs. (2) 52 mm. long, 32 pairs of legs. (3) 13 mm. long, 

31 pairs of legs. 

Range. — Brazil (Santarem); Venezuela (Merida); Panama (San 
Pablo, Rio Tapia) ; Canal Zone (Barro Colorado Island, Balboa, Las 
Cascadas, Fort Sherman, Rio Chinilla). 

Notes. — Individuals from Panama and the Canal Zone represent the 
variety vagans Brues. 

On Barro Colorado island this species was usually found under 
fallen logs in the forest and in leaf mold, occurring also between the 
broad sheaths of old fronds and the truniv of several of the large palms. 
It was met with occasionally in the ground-termite nests where it was 
discovered when the test stakes were pulled up. 

EPIPERIPATUS EDWARDSII (Blanchard) 

Peripatus edwardsii Blanchard, Ann. Sci. Nat., zooL, ser. 3, vol. 8, p. 140, 1847 
(Cayenne) ; Rech. Anat. et Zool. faites pendant un Voyage en Sicile, pt. 3, 
p. 64, 1849. 

Peripatus edwardsi Bouvier, Ann. Sci. Nat., zool., ser. 9, vol. 2, p. 301, pi. 9, 
figs. 74-79, text figs. 5, p. 15, and 111, p. 308, 1905 (Panama; Dari6n).— 
Fuhrmann, M6m. Soc. Neuchat. Sci. Nat., vol. 5, p. 190, 1912 (Brazil and 
Guiana to Panama). 

Epiperipatus edwardsi A. H. Clark, Proc. Biol. Soc. Washington, vol. 26, p. 18, 
1913 (listed); Smithsonian Misc. Coll., vol. 63, No. 2, p. 2, 1914 (Isthmus 
of Panama; from Bouvier); vol. 65, No. 1, p. 23, 1915 (Panama; Dari6n; 
from Bouvier). — Fuhrmann, Abh. Senck. naturf. Ges., vol. 36, Heft 2, pp. 
277-283, 1915 (listed). 



OXYCHOPHORES OF PANAMA — CLARK AND ZETEK 211 

New Records. — El Cermcno, Panama; J. Zctek, Jiily 8, 1941. 
Female: (O 35 mm. lonp;, 2 mm. broad, 30 pairs of logs. 

Balboa; J. Zotek, 1944. Females: (1) 50 mm. long, 34 pairs of legs. 
(2) 50 mm. long, 33 pairs of logs. (3) 45 mm. long, 33 pairs of logs. 
(4) 35 mm. long, 33 pairs of legs. (5) 25 mm. long, 33 pairs of logs. 
Males: (1) 28 mm. long, 29 pairs of legs. (2) 27 mm. long, 29 pairs 
of legs. (3) 25 mm. long, 30 pairs of logs. (4) 25 mm. long, 29 
pairs of legs. (5) 25 mm. long, 29 pairs of legs. 

Range. — Caj'enne; Surinam (Paramaribo); Venezuela (Caracas, 
M6rida, Valencia, Haut-Sararc, Bas Sarare); Colombia (Santa 
Marta); Panama (El Cermeno, Panama Station, Panama Railway); 
Canal Zone (Balboa); Dari6n. 

Notes. — At El Cermeno this species was found in situations similar 
to tliose frequented by Oroperijyaius corradi, though more often under 
boaj-ds resting on the ground. 

EPIPERIPATUS BIOLLETI (Boavier) 

Peripalus biolleyi Bouvier, Bull. Soc. Ent. France, 1902, p. 258 (San Jos6, Costa 
Rica) ; Ann. Sci. Nat., ser. 9, zooi., vol. 2, p. 321, pi. 10, fig. 85, text figs. 115, 
116. p. 323, and figs. 117, 118, p. 324, 1905. 

Epiperipalus biolleyi A. H. Clark, Proc. Biol. Soc. Washington, vol. 26, p. 18, 
1913.— FuHUMANN, Abh. Senck. naturf. Gcs., vol. 36, Heft 2, pp. 277-283, 
1915. — A. H. Clark, Proc. U. S. Nat, Mus., vol. 85, p. 3, 1937 (Parismina 
and La Caja, San Jos6, Costa Rica). 

New record. — El Cermino, Panama; J. Zctek, July 1941. Females: 
(1)41 mm. long, 4 mm. broad, 30 pairs of legs. (2) 38 mm. long, 
3 mm. broad, 30 pairs of legs. 

Range. — Costa Kica (San Jos6, and La Caja, San Jos6, Surubres 
near San Mateo, and Parismina); Panama (El Cermeno). 

Note. — At El Cermeno this species occurred in much the same 
situations as E. edwardsii. 

ADDENDUM 

Dr. Otto Fuhrmann (1912, p. 190) gives Epiperipatus simoni 
(Bouvior) as ranging from Bnizil and Guiana to Panama. There are 
no published records of this species from anywhere west of Veno/iu la. 
He also included "Brazil" in the range of Epiperipatus edwardsii, but 
we know of no record of this species from that country. 



212 PROCEEDINGS OF THE NATIONAL MUSEUM voL.g» 

BIBLIOGRAPHY 

Blanchard, Emile. 

1847. Malacopodes (Malacopoda de Blainville). Ann. Sci. Nat., ser. 3, 

vol. 8, pp. 137-141. 
1849. Recherches sur I'organisation des vers. Pt. 3 of Milne-Edwards's 

"Recherches Anatomiques et Zoologiques Faites pendaot un Voyage 

sur las Cotes de la Sicile. . . ." 353 pp., 26 col. pis. 
BouviER, Eugene L. 

1899a. Sur les variations et les groupements sp^cifiques des P^ripates 

am^ricains. Comptes Rendus Acad. Sci. Paris, vol. 128, pp. 

1344-1346. 
1899b. Nouvelles observations sur les Peripates americains. Comptes 

Rendus Acad. Sci. Paris, vol. 129, pp. 1029-1031. 
1900. Observations nouvelles sur les Peripatus [Onych.]. Bull. Soc. Ent. 

France, 1900, pp. 394-395. 
1902. Peripatus biolleyi, onychophore nouveau de Costa-Rica. Bull. Soc. 

Ent. France, 1902, pp. 258-259. 

1905. Monographie des on3^chophores. Ann. Sci. Nat., ser. 9, vol. 2, Nos. 

1-3, pp. 1-383, 140 figs., 13 pis. 

1906. Observation biologique. Le Peripatus edwardsi au Br^sil. Bull. Soc. 

Ent. France, 1906, p. 268. 

1907. Catalogue des onychophores des collections du Museum d'Histoire 

Naturelle de Paris. Bull. Mus. Hist. Nat Paris, 1907, pp. 518-521. 

1908. Sur le Peripatus brasiliensis Bouv. BuU. Soc. Philomat., ser. 9, vol. 

10, pp. 50-52. 
1928. A propos des observations du Fr. Claude-Joseph sur un P^ripate du 

Chili. Ann. Sci. Nat., ser. 10, vol. 11, fasc. 2, p. 260. 
Brues, Charles Thomas. 

1911. A new species of Peripatus from Grenada, with observations on other 

species of the genus. Bull. Mus. Comp. Zool., vol. 54, pp. 306-318, 

2 pis. 

1913. Preliminary descriptions of two new forms of Peripatus from Haiti. 

Bull. Mus. Comp. Zool., vol. 54, pp. 519-521. 

1914. A new Peripatus from Colombia. Bull. Mus. Comp. Zool., vol. 58, 

pp. 375-382, 2 pis. 
1917. A new species of Peripatus from the mountains of northern Peru. 

Bull. Mus. Comp. Zool., vol. 61, pp. 383-387, 1 pi. 
1923. The geographical distribution of the Onychophora. Amer. Nat., vol. 

57, pp. 210-217. 
1925. Notes on Neotropical Onychophora. Psyche, vol. 32, pp. 159-165. 
1935. Varietal forms of Peripatus from Haiti. Psyche, vol. 42, pp. 58-62. 
Brues, C. T., and Melander, Axel Leonard. 

1932. Classification of insects: A key to the known families of insects and 

other terrestrial arthropods. Bull. Mus. Comp. Zool., vol. 73, 

672 pp. 

(Onychophora, pp. 532-533, figs. 998-1002.) 
Camerano, Lorenzo. 

1898. Viaggio del Dr. Enrico Festa nella Repubblica dell'Ecuador e region! 

vicini, VII: Onicofori. Bull. Mus. Zool. Anat. Comp. Univ. Torino, 

vol. 13, No. 316, pp. 1-3. 



ONYCHOPHORES OF PANAMA CLARK AND ZETEK 213 

Clark, Austin Hobart. 

1913a. A revision of the American species of Peripatus. Proc. Biol. Soc. 

Washington, vol. 26, pp. 15-20, Jan. 18. 
1913b. Notes on American species of Peripatus, with a list of known forms. 

Smithsonian Misc. Coll., vol. 60, No. 17, pp. 1-5, Jan. 25. 
1913c. Piccolc note su degli Onychophora. Zool. Anz., vol. 42, pp. 253-255, 

July 18. 
1914a. Notes on some specimens of a species of onvchophore {Oroperipalus 

corradoi) new to the fauna of Panama. Smithsonian Misc. Coll., 

vol. 63, No. 2, pp. 1-2. Feb. 21. 
1914b. On some onychophores {Peripatus) from the Republic of Panama. 

Zool. Anz., vol. 45, pp. 145-146, Dec. 4. 
1915a. The present distribution of the Onychophora, a group of terrestrial 

invertebrates. Sniithsonian Misc. Coll., vol. 65, No. 1, pp. 1-25, 

Jan. 4. 
1915b. A note on the occurrence of Epiperipatus imthurmi (Sclater). Proc. 

Biol. Soc. Washington, vol. 28, p. 182, Nov. 29. 
1929. Peripatus from the Island of Montserrat. Proc. Ent. Soc. Washing- 
ton, vol. 31, p. 139. 
1937. On some onychophores from the West Indies and Central America. 

Proc. U. S. Nat. Mas., vol. 85, pp. 1-3. 

COCKERELL, ThEODORE D. A. 

1913. A Peripatus from Guatemala. Proc. Biol. Soc. Washington, vol. 26, 
pp. 87-88. 
FuHRMANN, Otto. 

1912. Quelques nouvcaux Pdripates am^ricains. M6m. Soc. Neuchdt. Sci. 

Nat., vol. 5, pp. 176-192, 16 figs. 

1913. t)ber einige neotropische Peripatus- Arten. Zool. Anz., vol. 42, pp. 

241-248. 
1915. Uber eine neue Peripatus- A.Tt vom Oberlauf des Amazonas. Abh. 

Senck. naturf. Ges., vol. 36, pp. 277-283, 1 fig., 1 pi. 
Mabcus, Ernst. 

1937. Un onychophoro novo, Peripatus (Epiperipatus) cvrlinae, sp. nov., de 

Goyaz. Rev. Mus. Paulista, vol. 21, pp. 903-910, 2 pis. 

PiCADO, C. 

1911. Sur un habitat nouveau des Peripatus. Bull. Mus. Hist. Nat. Paris, 
1911, pp. 415-416. 
Wheei.er, William Morton. 

1898. A new Peripatus from Mexico. Journ. Mon^h., vol. 15, pp. 1-8, 1 pi. 



■. t. •ovimaiNT mrRTiii* orricit if4« 



PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 

issued 




SMITHSONIAN INSTITUTION 

U. S. NATIONAL MUSEUM 



Vol. % Wathington: 1946 Nq. 3198 

ECHiraOID AVORMS OF THE NORTH PACIFIC OCEAN 



By Walter Kenrick Fisher 



The echiuroids, sometimes called spoon worms from the shape of the 
contracted prostomium or proboscis, are cigar-shaped or sausage- 
shaped creatures, essentially highly muscular sacs filled with fluid in 
which the long alimentary canal and other organs have great freedom 
of movement. The mouth is anterior, usually at the base of a snout 
or a long proboscis used for gathering food. The skin is highly gland- 
ular and is covered by a very thin cuticle. Typically there are two 
hooked setae behind the mouth, and two genera have either one or two 
circles of setae at the posterior end of the body. The alimentary canal, 
in contrast to that of most annelids, is several times the length of body 
and consists of a long foregut, differc'ntiated into pharyiLX, esoi)hagus, 
giz/.ard, and stomacii, a still longer midgut or intestine, accompanied 
for a considerable part of its length by a collateral intestine, or siphon, 
)ind finally a short hind-gut or cloaca, into which empty two usually 
\<)hiniinc)us, sometimes branched, vesicles, the walls of which are 
studded with minute ciliated funnels. The anterior nephridia, 
tyj)ically elongate, thin-walled sacs, varying from one to many, but 
usually from one to four pairs, hav(! a basal or terminal nephrostome, 
the lips of which may be greatly prolonged and spirally twisted. The 
nephridia vary greatly in size and when filbnl with eggs or sperm ar(^ 
(tftcn very large. The vascular system consists of a ventral vessel 
f(»llowing the nerve cord to form a looj) in the proboscis from the tip of 
u hich a median vessel pa.sses backward in tiie proboscis and along the 
dorsal side of foregut to beginning of midgut, where a neurointestiiuil 
vessel joins it to the ventral vessel. In Urechis there are no blood 
vessels. 

215 
•""oaa*— *fl — 1 



216 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

Echiuroids are burrowers in mud or sand, where they fashion more 
or less permanent tunnels. Sometimes they live under rocks; some- 
times in mud-filled mollusk shells or sand-dollar tests, which afford 
some protection; or they inhabit the rock galleries excavated by boring 
clams. Their food consists of organic material contained in the mud 
which they swallow in large quantities, or of lighter organic detritus 
selected by the usually long proboscis. In the same species from dif- 
ferent locaUties the intestinal pellets vary with the character of the 
bottom. One genus (Urechis) has very specialized feeding habits and 
uses only finely divided material, including bacteria. It is probable 
that any small organisms living in the surface film of mud will be 
eaten by echiuroids. Gislen (1940, p. 30) found the intestinal pellets 
of Echiurus echiurus "to consist of the same stuff as that which is 
formed by the detritus-film growing on the aquarium bottom. There 
are thus plenty of sand grains; further the pellets consist of diatoms, 
algal threads, debris of leaves of phanerogams (Zostera et al.), 
Infusoria, Bacteria, occasional Nematodes and Rotatoria and, to a 
large extent, of amorphous brown stuff which emanates from 
decomposed organic substance." 

The smallest sexually mature echiuroid I have ever seen is a 
Listriolobus pelodes 7 mm. long (0.275 of an inch), and the largest is 
Urechis caupo, 470 mm. long preserved, or 18.5 inches. I have seen 
a relaxed living Urechis caupo 19.75 inches long. The Japanese 
Ikeda taenioides, a remarkable and isolated form, attains a body 
length of 16 inches with a proboscis of 58 inches, or a total length of 
6 feet 2 inches (Ikeda, 1907, p. 20). 

More helpless, unprotected animals can scarcely be imagined. The 
immature stages are prey for every predaceous inhabitant of the 
sea bottom. The adults are regularly eaten by fishes, especially 
flatfishes and rays, as well as by the Indians of Chiloe Island, Chile 
(Gay, 1854, p. 475). In Japan and Korea Urechis unicinctus is 
extensively used as bait. Sato (1939, p. 319) states that in Korea 
the natives catch it by means of iron hooks and dry it for food. 

Systematics of the Echiuroidea present the usual problems in 
addition to others inherent to the group. The principal diSiculty is 
the lack of structures having a permanent form. The setae are of 
very limited use; everything else is soft and capable of distortion. 
The practical difficulties encountered are those which would confront 
the student of holothurians if these creatures did not carry embedded 
in the skin a species label in the guise of characteristic calcareous 
deposits. Most of the generic and specific characters of echiuroids 
must be sought by careful dissection of the internal organs, which 
arc susceptible to variation arising from accidents of fixation. Never- 
theless, a fairly satisfactory system of genera can be constructed. 
But it is obvious that closely related species may not be recognized, 



ECHIUnOID WORMS OF NORTH PACIFIC — FISHER 217 

or if recognized they may be impossible to describe in the absence 
of trenchant characters. Descriptions of echiuroids based on external 
charactei-s or on a very summary enumeration of a few internal 
features have made it difhcult to determine the generic position of a 
number of described forms. 

It has been a time-honored procedure to classify the echiuroids, 
sometimes in combination with the sipunculoids and priapuloids, 
under the name Gephyrea, as a class of the Annelida. In 1898 Prof. 
Adam Sedgwick, in his "Students' Te.xtbook of Zoology," set up 
separate phyla for the Sipunculoidea and Priapuloidea but retained 
the Echiuroidea as a class of the Annelida. Since the development 
of I rcchis caupo has been thoroughly elucidated (Xewby, 1940), 
it is now known that the echiuroids are not more closely related to 
annelids than to moUusks. Dr. Newby writes at length on a com- 
parison of echiuroid development with that of the other invertebrates 
and on the phylogenetic position of the Echiuroidea. In conclusion 
he says (p. 209): 

There are many echmroid characteristics which indicate that this group is 
separate from the annelids: (a) The mode of development of the first somatoblast 
is different, (b) The anus is not homologous in the two groups and no procto- 
daeum is formed in echiuroids. (c) The mesodermal bands do not develop 
teloblastically in echiuroids. (d) The elongation of the larva is not teloblastic 
in echiuroids. (e) Three laj'ers of body muscles are formed in echiuroids. (f) 
The ectomesoderm contributes to the body musculature in echiuroids. (g) A 
ciliated intestinal groove is formed in echiuroids and this becomes the primordium 
of the siphon. These structures are not found in the annelids, (h) Anal vesicles, 
probably of endodermal origin, are found in echiuroids. (i) The coecuin of the 
larval digestive tract becomes a linear part of the adult tract in echiuroids. (j) 
The mesodermal bands of echiuroids show no evidence of segmentation, (k) 
The lack of segmentation in the mesoderm considered with the questionable 
nature of the segmentation of the nervous system and mucous glands and further 
considered with the "segmentation" of the shell glands of the chitons (molluscs) 
makirs it appear probable that the echiuroids have a i^riinary lack of metamerism. 

.\gainst these numerous difTercnccs there are only three clear-cut characteristics 
in common between the echiuroid.s and annelids which are not also po.ssessed by 
the mollusks. (a) The annelidan cro.ss develops in both groups, (b) Both 
groups possess setae. (However setae are to lie found in another group of animals 
(Brachiopoda) which do not belong to the Annelida.) (c) The lateral halves of tho 
nervous system become merged into single, unpaired structures. 

With the above facts in mind it is evident that the echiuroids are only distantly 
related to the annelids. When numerous differences which appear in their 
fievelopment are con.sidcrerl, it seems improbable that the inclusion of the echiu- 
roids with the annelids as a sub-phylum or cla.ss, is justified. It is probably more 
accurate to consider the Echiuroidea as forming a separate phylum, di.stinct from 
the f)hylum Annelida, and I herewith i)roi)()se that they be so Cftusidered. 

In the keys no nicntion is made of Kpithetosoma Danielssen and 
Koren, 1881. Th6cl (1900, p. 9) has demonstrated satisfactorily 
that the animal is not an echiuroid, but most likely a nemertean. 



218 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 96 

Neither is Poeobius meseres Heath (1930) included. This remarkable 
pelagic transparent worm was first taken in 350 meters, Monterey 
Bay, Calif., and was later found to be abundant off southern Alaska. 
Its anatomy has been fully described by Professor Heath. Sub- 
sequently the writer observed and sketched a living animal. The 
blood vessels are clearly visible and contain a dull green fluid, but the 
enlargement of the dorsal vessel is dull red. Blood vessels extend to 
tip of the 2 prostomial palps and the 10 (possibly peristomial) cirri. 

The creature has no paired appendages, no somatic segmentation, 
and no setae. "The nervous system conforms to the usual annelidan 
type, with supra-oesophageal ganglion, circum-oesophageal connec- 
tives, and a ventral nerve chain comprising 11 pairs of ganglia with 
the usual commissures and connectives. The somatic musculature 
comprises four great longitudinal bands, extending throughout the 
length of the body, and a more delicate external sheath of circular 
fibres." This is the annelid pattern and distinctly not the echiuroid. 
The head is unlike that of any known echiuroid but resembles 
that of some polychaete annelids. The alimentary canal and 
nephridia seem to the writer to be specialized m much the same way as 
in the case of Sternaspis, which in one species {S. spinosus Sluiter) has 
the prostomium prolonged outward on each side to form a grooved 
palplike organ. The Scoleciformia, however, have definite mesoder- 
mal segmentation. 

The difficulty in finding a place for Poeobius may well mean that it 
is not an annelid or a echiuroid or a link between the two. Although 
nothing whatever is known of the development of Poeobius, we have 
to assume that mesodermal segmentation is absent; therefore it is not 
an annelid. Its nerve cord is segmented (implying pseudometamer- 
ism). The nerve cord of larval echiuroids is segmented, but this is 
lost in the adult, suggesting that the ancestors, while deprived of 
mesodermal metamerism, still had a pseudometamerism of the nerve 
cord. A tenable hypothesis is that the echiuroids and Poeobius 
stemmed from a common group that was as fundamentally un- 
segmented as the Amphineura among mollusks. According to this 
view Poeobius is the survivor of a lesser phylum, comparable to the 
Phoronidea and Priapuloidea. As the genus now floats in a sort of 
taxonomic limbo, it may be provisionally assigned to a new phylum, 

POEOBIOIDEA. 

The region covered by this report includes all the water north of a 
line drawn from Cape San Lucas, Baja California, to the southern 
end of Sakhalin Island on the east Asiatic coast. The Gulf of Cali- 
fornia has been included, and a species long ago dredged by the 
Albatross in Japanese waters has been added, as it modifies the 
concept of Acanthohamingia, which I wished to include in the key. 



ECllILKOlU WUK.MS OF NUHllI I'AflFlC — FISHER 219 

Tlio spcciinons upon svhirh tliis ])apfr is based have been accuinu- 
lati'il slowly over a considerable perioti of 3'ears.' In addition, the 
material belonjrinf: to the United States National Museum was 
placed at my disposal, and an iiu])ortaTit collection belon<rin<2: to the 
Allan Hancock Foundation of the I'niviTsit}' of Southern California 
was tcndereil by Dr. C)l<;a Ilartman. The types of all the new species 
are in the collection of the National Museum. 

The followino; ni'W <^eiuis, based on an extralimital species, will be 
found in the text: Lissomyema, tyj)e Thalassema mellita Conn (under 
Listriolobus). 

Phylum Echiuroidea' 

Ik'hiuroidea Sedgwick, 1898, p. 527 (class of Annelida). 

Unsegmented. bilateral, fusiform or saccuUform animals with 
anterior mouth and posterior anus, but no proctodacum; a long con- 
voluted alimentary canal lying in a spacious coelom of scliizocoelous 
type; a muscular body wall composed of three layers, of which the 
middle (with one I'xception) is composed of longitudinal fibers; with 
one to very numerous anterior nephridia functioning as gonothecae; 
with typically two anal vesicles having numerous ciliated fuimels 
and functioning as excretor}' organs; alimentary canal tyjiically with 
collateral intestine or siphon; usually with a ])rostomial proboscis, 
which nuiy exceed length of boily but which is sometimes absent; 
usually with ectodermal setae, of which two, ventrally situated be- 
hind mouth, are most constantly present, together with sometimes 
one or two circles at posterior end of body; but setae absent in a few 
genera; ventral nerve cord unsegmented forming, around the mouth, 
a loop which follows border of proboscis; gonad, where known, in 
mesentery above nerve cord, or hi the mesenteries surrounding cloaca. 

KEY TO CLASSES 

o'. Body wall with inncniiowt circular or oljliijiic layer uf inu«ck's well dcvoloped; 
anal vcHicles present; collateral intestine or siphon well developed; pro- 
bosciH and anterior setae present in nearly all species Echiurida (p. 220) 



• I am Mix-cifilly ln-holflon to niy foriiuT colloiKiic I'rof. fii'<)rKe E. Mncniiilllc for iiiiitcriMi of I'rrrhit 
caupo, LiMriolobu* prlodfM, nrul Ochdottoma ocloinvotum; to Eilwnrd F. Kickctts for tlu' type of l-xhiurua 
eehiurui ataikanut nn<l n sriinll colli-ctlon miiric by lilni and Join) Stcliilx'rk In the (liilfof C'liliforniii: to Dr. 
Olitn llHrtiiinn (or n spccltncii of Limnmvrmn mrlliln: to I'rof. S. K. Lluht for ft [M-rfcct sihtImicii of lAntrMnlitu 
ptlodtt; Ui tlm Mii.wum of Companitivo '/ool'>Ky for a .^in'i-lmoii of L'reckin chtlentin; to Dr. \V. I,. I.loyd, 
Cnbrillo .Vlarlni^ Miiacum, San I'wlro, Calif., for the loan of a spiclnii'ii of Ochiloitoma octorni/ntum; to I'rof. 
John n. (iprnuM for the loan of scvcml rare rcprlnU; and to Dr. Waldo L. Schinitt, U. S. National Museum, 
for numerous favors. 

• Kchluroldra wa.i Introdutwl m a subphylum by A. II. riark (Hull. Inst. OcAannpr. Monaco, No. 400, 
p. 24, 1021) and a.s a [iliyluni by W. W. Newby (IWO. p. 210) Bn<l I.lbblc II. Ilyinan (HMO, pp. 34, M). As a 
matter of record, F.dwaril K. Hlcketls wiwt the first to u.ifl Kcliluroldi-a aa n i>liyluin name. In an exwllont 
semlpopulnr text "Betweon Pacific Tl«|t«" (UlcketLs and C'ajvln, 19.10, p. 272), i>ertiap.s incited thereto by 
tho present writer who has a>l%'ocnted this proc«<luro to his classes for 20 ycnrs. 



220 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

0^. Body wall with innermost circular layer missing or degenerated to a net 
of fibers; no anal vesicles; apparently no siphon; no proboscis and no 
setae Sactosomatida » 

Class ECHIURIDA 

KEY TO ORDERS 

o'. In body wall longitudinal muscle layer lying between outer circular layer 
and inner oblique layer; nephridia, normally paired, not excessively nu- 
merous. 

6*. A closed blood- vascular system; no specialization of intestine for anal 

respiration Echiuroinea Bock (p. 220) 

62. No vascular system, coelomic fluid being heavily charged with large blood 
corpuscles containing hemoglobin or hemoglobin plus hematin; in- 
testine with terminal portion enlarged, thin-walled, to receive water 

from cloacal pump Xenopneusta, new order (p. 262) 

a?. Longitudinal layer of body wall lying outside of both the circular layer and 
inner obHque layer; nephridia excessively numerous, unpaired (and with 
terminal nephrostome) ; proboscis excessively long. 

Heteromyota,* new order 

Order Echiuroinea Bock, emended 

A closed blood-vascular system; no specialization of intestine for 
anal respiration. 

KEY TO FAMILIES 

ci. Dimorphic; male degenerate, planarianlike, parasitic in or on female; female 
resembling Thalassema but with bifid proboscis in some genera; anal vesicles 
consisting of branched tubules ending in numerous cihated cups; anterior 
setae sometimes present; posterior setae absent Bonelliidae (p. 249) 

a?. Not dimorphic; proboscis usually conspicuous, sometimes several times length 
of body, but never bifid; absent in one genus; anal vesicles not branched 
but in form of elongate sacs, surface of which is covered with minute cilated 
funnels; anterior paired setae present in all genera, posterior setae in 
Echiurus only Echiuridae (p. 221) 



3 New name for Saccosomatida Thfiel (1906, p. 14). Th§el instituted the group as a suborder for Sacco- 
soma vitreum Danielssen and Koren (1881, p. 34, pi. 6, figs. 1-8). This species is based on a single small 
example dredged in 1,215 fathoms north of the Faroe Islands. It is a female and the species may prove to 
be dimorphic, as there is a single nephridium filled with eggs and opening near the mouth. The proboscis 
may have been lost. It is aberrant from all other echiuroids and may not be an echiuroid. Saccosoma 
Danielssen and Koren is preoccupied by Saccosoma Motschoulsify, 1859, in Coleoptera (Bull. Acad. St. 
Pfetersbourg, vol. 1, column 304). The new name "Sactosoma" (with identical meaning) is proposed to 
replace Saccosoma Danielssen and Koren. 

* Based on the remarkable genus Ikeda Wharton, 1913, pp. 243-270. Type, Thalassema taenioides Ikeda, 
1904, p. 63; 1907, p. 16, pi. 1, fig. 3; pi. 2, figs. 18-22; pi. 3, figs 23-36; pi. 4, figs. 37-47. This large echiuroid, 
with a proboscis upward of a meter or more in length and nephridia from 200 to 400 in number without 
indication of paired arrangement, is so different from the general run of the phylum that it deserves to be 
set apart as the type of at least a distinct order. The arrangement of muscle layers is different from that of 
all other echiuroids and indicates a long separation from typical stock. See Sato, 1931. p. 179. 



ECHIUROID WORMS OF NORTH PACIFIC — FISHER 221 

Family ECHIURIDAE (de Blainville, 1827, restricted) 

KEY TO GENERA 

a'. Two circles of posterior setae Echiurus Gu6rin-M6ncville (p. 225) 

a*. No posterior setae present. 

b^. Proboscis absent Arhynchite Sjito (p. 247) 

6*. Proboscis present. 

c*. No differentiated thicker bands in longitudinal muscle layer, 
d*. Nephrostome of nephridia without elongated, si)irally coiled lips. 

Thalassema Lamarck (p. 230) 
(P. Nephrostome with elongated, spirally coiled lii)s. 

Anelassorhjmchus Annandale (p. 221) 

c*. Longitudinal muscle layer with very slight to pronounced differentiation 

into longitudinal bands, 8 or more in number. 

d'. Nephrostome of nei)hridia without spirally coiled lips; inner layer of 

muscles not differentiated into separate transverse fascicles between 

longitudinal bands Lissomyema, new genus (p. 224) 

(P. Nephrostome with elongated spiral lips. 

e'. Differentiated longitudinal muscle bands weak, zones between not 
showing a fasciculate arrangement of inner oblique muscles; in 
small specimens longitudinal bands very faint or visible only in 

posterior region Listriolobus W. Fischer (p. 233) 

e'. Longitudinal muscle bands strongly developed, zones between 
crossed by separated fascicles of innermost, oblique layer. 
p. Nephridia in 1 to 5 pairs; vascular ring vessel at beginning of 

midgut Ochetostoma Lcuckart and Ruppell (p. 240) 

p. Nephridia, at least in male, in to 14 groups of 1 to 4, the groups 
arranged in pairs; vascular ring vessel at posterior end of 
pharynx Ikedosoma Bock (p. 224) 

Remarks. — In the foregoing synopsis all the generic divisions, 
with the exception of Echiurus, are the result of subdividing the old 
genus Thalassema. In a very real sense these groups are provisional 
because adequate descriptions and figures of the internal structure 
of many species have not been published. 

Thalassema Lamarck.^ — The geiuis has been restricted to a few 
species grouped around the type, Thalassema thalassema (Pallas), 
generally known as lli. neptuni Gaertner. The middle, longitudinal 
layer of muscle fibers of body wall shows no sign of differentiation 
into thicker bands. The internal opening of the nephridia is very 
simple, without prolongation into si)irally coiled lips. 

ANELASsoRHYNrHis Aiuiandale (1922, p. 148). — It may not be of 
any practical value to recognize this group. The species dilfer from 
Thalassema in having the nephrostome lips j)rolonged and spirally 
coiled, but little is known of other details of the internal anatomy. 
Amiundalc based the genua on the structure of the |)r()boscis of four 
cstuarinc species occiiniiig in lnackish water of India and Siam. 
He says: 

The genus con.sists of Kchiuridae allied to ThnlnHmma Gaertner, but differing 
in the structure, function, and phy.siology of tlic proboscis. Thi.s organ is rela- 
tively stout and short, incapable of great prolongation or autotomy. The ciliated 



222 PROCEEDINGS OF THE NATIONAL MUSEUM vol. se 

groove on its ventral surface is feebly developed and the lateral margins of the 
ventral surface bear (except in A. microrhynchus) gill-like outgrowths. The 
longitudinal muscle-fibres of the body form a single sheath and the musculature 
bears a close resemblance to that of some species of Thalassema. There are two 
pairs of nephridia. The anal funnels are simple and thin- walled; their ciliated 
funnels are minute. 

The type-species is A. branchiorhynchus (Annandale & Kemp). The other 
species are A. dendrorhynchus (Annandale & Kemp), A. sabinum (Lanchester) 
and A. microrhynchus (Prashad). 

It seems to me that the modifications of the proboscis, which 
exhibit a number of gradations in complexity, are adaptations to an 
ecology in various ways abnormal, a parallel development being 
found in Ochetostoma arkati (Prashad). But these species agree with 
certain others in having a more specialized nephi-ostome than is 
found in Thalassema thalassema and close allies. 

1. With two pairs of nephridia (behind the setae): sabinum Lanchester, 
branchiorhynchus Annandale and Kemp, dendrorhynchus Annandale and Kemp, 
microrhynchus Prashad, semoni Fischer. 

2. With three pairs of nephridia, all three opening behind the setae: mucosa 
Ikeda, vegrande Lampert (no proboscis). First pair opening in front of setae: 
inanense Ikeda, moebii Greef. 

Unless some definite character other than the nephrostome is dis- 
covered, there will be a practical difficulty in distinguishing young 
Listriolobus , in which the differentiation of longitudinal muscle 
bands is very weak. 

LissoMYEMA. — Through the kindness of Dr. Olga Hartman I 
have received a specimen of Thalassema mellita Conn collected by 
her at the type locality, Beaufort, N. C, in June 1940. It is 36 
mm. in length, with proboscis 16 mm. additional. From the outside 
the eight longitudinal muscle bands are clearly visible. Figure 10 
represents a dissection of the anterior portion. The muscle bands 
are much more sharply delimited than in Listriolobus by having an 
incipient fasciculation of the muscles of the oblique layer, possibly 
representing the first stage in the differentiation of the strong trans- 
verse bundles characteristic of Ochetostoma. The species has simple 
fan-shaped nephrostomes and very heavy interbasal and radiating 
seta muscles. The gizzard is relatively short and the stomach (C) 
is relatively long. An individual variation is the presence of three 
nephridia on one side and two on the other. The species is described 



Figure 10.^ — Lissomyema mellita (Conn): Dissection of anterior region of a specimen from 
Beaufort, N. C, X 12. Six of the eight muscle bands are diagrammatically indicated by 
dots. The alimentary canal is drawn to the right to disclose the organs beneath it. 
{B^, &, B*, dorsal, neurointestinal, and ventral blood vessels; C, stomach; G, gizzard; 
/, intestine; MD, middorsal muscle band; MI, interbasal muscle; MF, midventral muscle 
band; N, nephridia; NC, nerve cord; O, esophagus; P, pharynx; S, seta; Sil, beginning of 
siphon.) 



ECHIUHOII) W01{MS OF XOHTH PACIFIC — FISHER 223 




FiGURK 10.— (Sec opposite page for legend). 



224 PROCEEDINGS OF THE NATIONAL MUSEUM vol.98 

as having two pairs. The anal vesicles are voluminous with numerous 
conspicuous ciliated funnels. The ventral blood vessel sends an 
important branch to the pharynx and esophagus. 

For this species, therefore, I propose the new genus Lissomyema, 
which differs from Thalassema in having eight well-differentiated longi- 
tudinal muscle bands and incipient fasciculation of the oblique layer; 
from Listriolobus and Ochetostoma in having simple fan-shaped nephro- 
stome without trace of spiral extensions. Type, Thalassema mellita 
Conn. (Fig. 10.) 

Ikedosoma Bock. — Thalassema elegans Ikeda^ does not belong in 
Thalassema. Ikeda (1907, p. 50) writes: "All the longitudinal lines 
visible on the outside, excepting the one which runs in the mid-ventral 
line and is superposed by the nerve-cord, appear on the inner surface of 
the body-wall as slightly elevated narrow ridges or thickenings oj the 
longitudinal muscular layer. In the ten zones separated from one 
another by the above lines, the circular muscle fibres form more or 
less regularly arranged transverse bundles." This structure of the 
body wall closely approximates that of Ochetostoma, but elegans is 
peculiar in having numerous (13 to 27) nephridia in six or seven 
pairs of groups comprising one to three nephridia each. "The internal 
opening present at base is provided with 2 relatively short spiral 
lobes." The dorsal blood vessel ends with the "heart" on the hind 
end of the pharynx and is therefore shorter than in typical Echiuridae. 
"The neuro-intestinal vessel arises from the ventral median point of 
the ring-sinus, which surrounds the extreme hind end of pharynx" 
{ibid., p. 52). There is no interbasal muscle and no intestinal coecum. 

Thalassema gogoshimense Ikeda (1904, p. 66, pi. 1, fig. 19) is ap- 
parently congeneric with elegans. The excellent colored figure shows 
the same white longitudinal stripes as elegans, reflecting the muscular 
structure of body wall. Ikeda says: "It shows an essential agreement 
with Tlialassema elegans. Indeed, the agreement may be said to be 
complete, the only difference being that all the visceral organs in the 
present species are developed on a smaller scale in proportion to the 
smaller size of its body." In the females, however, the nephridia 
are present in three pairs all situated behind the setae, while in the 
male they are in six to eight groups, arranged in pairs, each group 
with one to four nephridia, which are like those of elegans in structure. 

The above paragraphs were written and a name was assigned to 
the genus before I saw Bock's paper. They are retained since we 
independently arrived at the same conclusion. 



« Ikeda, 1904, p. 66; 1907, p. 47, pi. 1, fig. 4; pi. 4, figs. 48, 49; Sato, 1939, p. 366; Bock, 1942, p. 18. 



ECHIUROID WORMS OF NORTH PACIFIC — FISHER 225 

Genus ECHIURUS Gu^rin-iMeneville 

Echiurus GifRiN-MfiNEViLLK, 1831, p. 9, pi. 6, fig. 3 (ex Shipley, 1899, p. 342) 
(type, Lumbricus echiurus TaUas, 1700= Echiurus pallasii CUi<5rin-M<5nevilIe, 
1831). — Skorikov, 1909, p. 80. — Spengkl, 1912b, p. 173. 

Echiuriclae with two rings of posterior bristles, a well-developed 
proboscis, two or four nepliridiii (without spirally coiled lips), and a 
postpharj-ncjeal tliaphragni, which separates incompletely the small 
liead coelom from the perivisceral cavity. 

ECHIURUS ECHIURUS ALASKANUS. new subspecies 

Plate 20 

Echiurus Pallasii C. B. Wilsov, 1900, p. 174. 
Echiurus echiurus Spengel, 1912b, p. 183. 

Diagnosis. — Differing from typical E. echiurus (Pallas) of the north 
Atlantic and neighboring Arctic Ocean in having the proboscis strongly 
attached to the body and in having the posterior setae definitely 
curved lather than nearly straight. Length of type, 230 mm. plus 
much contracted proboscis, 20 mm. 

Description. — ^Length of body upward of 230 mm., commonly 100 
mm., stout; proboscis adherent, fleshy, convex above, the edge in- 
curved ventrally, su})truncate distally, usually 15 to 20 mm. long in 
contracted state. On ventral suHace of the proboscis a difTcrentiated 
thickening extends as a low ridge from the mouth for about one- 
fourth length of proboscis but sometimes considerably farther. The 
integument is roughened by rings of prominent vermcae most crowded 
at ends of body. In the middle region, where they are generally 
loss crowded, rings of more prominent verrucac alternate with zones 
of three to five nngs in which the verrucac are smaller or more widely 
spaced, or both. The appearance depends largely upon the degree 
of contraction of the body muscles. 

The anterior setae are stout, strongly curved, and situated back of 
the base of proboscis a distance e([ual to about its greatest width. In 
each circle of anal bristles there may be variations of G to 8, as: 8-8 
(posterior ring); 8-7; 8-G; 7-7; 7-G. In some specimens where the 
number is less, inequality of spacing indicates loss of setae. These 
pf)stcrior setae \t\ry in degree of curvatun* but are slightly more 
curved than in typical A', ecldurus. The anterior setae have a strong 
interbasal muscle connecting their inner ends. Occasionally a second 
seta, in process of formation, a(;comi)anies one of the primary. The 
principal posterior muscle from the setae attaches to body wall just 
behind the anterior nephridium. 

The inner circular layr of body muscle shows n division into closely 
placed fascicles at each end of body, where the animal is nonnally 
most contracted. 



226 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

Nephridia 4, the anterior pair close behind the setae and a little 
farther from nerve cord. The funnel is conspicuous, with an undu- 
lating or frilled border, but is not prolonged into spiral lips. The 
nephridia of all specimens examined (taken in summer months) were 
contracted. In some cases they were very small, and the anterior 
pair very inconspicuous. 

Anal vesicles are simple, elongate, thin-walled sacs attached to 
ventrolateral wall of the cloaca and closely beset with minute ciliated 
funnels. 

Diaphragm (pi. 20, figs. 1, 4). The diaphragm is a curious, thin- 
walled, funnel-shaped septum incompletely separating the peripharyn- 
geal coelom from the general body cavity. Its general form is best 
appreciated from the figure in which it is shown in a semidiagrammatic 
fashion. The anterior, rouglily circular edge is completely attached to 
the body wall, while ventrally it is attached to body wall on each side 
of the nerve cord (which here lies within the ventral mesentery of 
pharynx and esophagus). A large oblique posterodorsal opening of the 
diaphragm (with complete free edge) allows the esophagus (with its 
strong ventral mesentery) to pass backward into the general coelom, 
sometimes above and sometimes below the interbasal muscle. The 
rim of aperture apparently has a sphincter. The two halves of the 
double ventral mesentery of esophagus merge with diaphragm along 
its paraneural part and a short distance above the nerve (pi. 20, fig. 5). 

Alimentary canal. The pharynx remains always in the peri- 
pharyngeal coelom. It is attached to the body wall by numerous strong 
muscular strands having an annular arrangement. There is a con- 
tinuation forward of the double ventral and dorsal mesenteries sep- 
arated into frenula. The dorsal blood vessel lies in this mesenterial 
complex. The head cavity is therefore much occluded by tissue. 
The lining of pharynx is anteriorly thrown into coarse folds. 

The esophagus begins just behind the region of the radiating frenula 
of pharynx. It has, in the anterior portion, a dorsal mesentery of 
slender separate strands, but there is a double membranous ventral 
mesentery throughout its whole extent. This mesentery is anchored 
in the peripharyngeal chamber on each side of the nerve cord, where, a 
short distance above body wall, it merges with the diaphi-agm. By 
means of its muscular mesenteries, all of the esophagus can be with- 
drawn mto the head cavity. 

The esophagus, on passing through the right side of the diaphragm 
close to posterior border, becomes a long gizzard, marked by rings, 
which are prominent amiular ridges of the lining. Beguming with the 
gizzard the alimentary canal is moored only by dorsal mesenterial 
ribbons as far as the cloaca, which has radiating muscular frenula. 
Along the dorsal side of the gizzard held by a perforated mesentery 



ECHIUROID WORMS OF NORTH PACIFIC — FISHER 227 

is the voluminous dorsal blood vess(>l with numerous papilliform 
l)ranchos, at least anteriorly. 

A very short rudimentary stomach or crop lies between the gizzard 
and liesritming of intestine (indicated by the ventral ciliated p-oove). 
The lining of stomach is thrown into 12 strong longitudinal folds, 
contrasting sharply with the annular folds of gizzard. Where tiie 
stomach becomes intestine, the dorsal blood vessel splits to form tiie 
ring vessel. 

The intestine has the usual three parts: presiphonal, siphonal, and 
postsiphonal. The first is about as long as the gizzard, or a little 
longer if relaxed. 

The siphonal part, rougldy 20 to 25 times length of presiphonal part, 
is marked by longitudinal folds of the lining which arc evident super- 
ficially. The siphon is about one-fourth the diameter of the intestines. 

The postsiphonal intestine has thinner walls and is about 10 times 
the length of presiphonal segment. The ciliated groove forms a 
ridge along its ventral side, and ends at a coecum (not always inflated) 
just in front of the cloaca. The fecal pellets which fill this part of the 
intestine are elongate ellipsoids and sometimes contain coarse material. 
I have found leaves of the hemlock (Tsuga). 

Vascular system. This consists of a dorsal and ventral blood 
vessel and neurointestinal connective. These vary in caliber in dif- 
ferent specimens. The dorsal vessel is likely to be considerably in- 
flated over part or the entire length of gizzard, vnih irregular lobose 
swellings anteriorly. The ventral vessel, attached to middorsal line 
of nerve cord, ends posteriorly as a solid cord just in front of the in- 
testinal coecum. The neurointestinal connective results from the 
brandling of the dorsal vessel at the beginning of intestine by which 
tlie neurointestinal ring (B^) is formed. The connective branches 
again (5^), to form the muscle ring, before merging broadly with 
the ventral vessel (B*). 

Type.— v. S.N M. No. 20G09. 

Type locality. — Auk Bay, Juneau, Alaska, collected by E. F. Rick- 
etts, August 14, 19:n. 

Specimens examined. — One hundred and twenty -four as follows: 

Collection of Stanford Univkk.sity • 

Kiikak Bay, Shelikof Strait, Alaska, 12 apccimcns, uruicr rocks, in iimd; McMillan, 

1924. 
\\ rangell, Ala.ska, 37 .si)ccimen.s; A. \V. Greely and R. K. Sno(lnra.ss, 1897. 
Auk Bay, near Juneau, Ala.ska, 3 specimens, slate beach, under rocks, in murldy 

.sand, lowest intertidal zone, July 17, 1931; K. V. Hickclts (also typo from 

this locality, August 14, 1931). 
Huston Inlet, Queen Charlotte Islands, Briti.sh ('(jlunihia, 42 specimen.^; W. F. 

Thornp.son, July 1, 1913. 
Alaska, pos.sibly Dutch Harbor, t'nalaska, 5 specimens. 



228 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

Collection of U. S. National Museum 

Cape Smyth, Alaska, 3 fathoms. No. 850, Point Barrow Expedition; 2 specimens 

in very bad condition, without proboscis ; possibly these are intermediate with 

true echiurus. 
Off Cape Strogonof, Alaska, Albatross station 3291, lat. 56°58'30" N., long. 

159° 11' W., 26 fathoms, black sand, gravel, 1 specimen from stomach of cod, 

in poor condition. 
Bristol Bay, Alaska, No. 4597, 2 specimens, one lacking proboscis. 
Unalaska, Alaska, No. 16314, 2 specimens. 
Bering Island, Nos. 4151, 16315, 2 specimens, typical. 
Alitak Bay (head of Lazy Bay), Kodiak Island, gravelly sand, January 22, 1941, 

5 specimens. 
Stepovak Bay, Alaska, 15-90 fathoms, October 24, 1940, 1 specimen. 
Dolgoi Harbor, Alaska, October 6, 1940, 1 specimen. 
Wrangell, Alaska, No. 4538, August 28, 1882, under stones, 2 specimens. 
Chasina Bay, Alaska, No. 4601, 1 specimen. 
Saginaw Bay, Alaska, No. 4117, 1 specimen. 
Without stated locality (probably Unalaska), 5 specimens. 

Remarks. — The type specimen was taken from its burrow, a tube 
in sandy clay beach, 24 inches below the surface, along with a com- 
mensal polynoid annelid, Hesperonoe adventor (determined by Dr. 
Olga Hartman). The same species is commensal with Urechis 
caupo. 

This, the common Alaskan Echiurus, differs from the typical form 
of Europe and the north Atlantic coast of America in having the 
proboscis firmly attached to the body. All writers who have handled 
living Echiurus echiurus emphasize its habit of dropping the proboscis 
on the slightest provocation.® Most of the 120 specimens of alaskanus 
were not handled with care but were simply dropped into alcohol; 106 
of these specimens still have the proboscis firmly attached, while at 
least 5 lost the proboscis subsequent to fixing, apparently from rough 
handling. 

Under ordinary circumstances I should have given Brandt's name 
sitkaensis to this form, assuming that Mertens would naturally have 
picked up at Sitka the common Alaskan species. J. W. Spengel 
(1912b), however, succeeded in obtaining one of Mertens's two spec- 
imens, upon which Brandt based his description, and found that it 
differed fundamentally from Echiurus echiurus in having only two 
nephridia, as well as in certain other respects. The name sitkaensis 
is therefore definitely associated with a type specimen which has been 
redescribed by one of the best zoologists of his time. 

The ecology of Echiurus echiurus has been studied by Dr. Torsten 
Gislen (1940) chiefly at Kristineberg, Sweden. His very compre- 



« Torsten Gislfin says: "As stated before the proboscis is very easily thrown off. In fact probably only 
very few men have seen a proboscis in connection with an Echiurus. Forbes and Qoodsir say that it is so 
slightly affixed to the body as to break off at the least touch; in only one or two cases did they find it attached, 
and then it broke away immediately on the removal of the animal. Only in some exceptional instances 
have I been able to secure specimens with the proboscis retained." (Gisl6n, 1940, p. 10.) 



ECHIUROID WORMS OF NORTH PACIFIC — FISHER 229 

hensive memoir is in the forefront of cxcollonco and will long servo as 
a model for this type of work. 

ECHIURUS SITKAENSIS (Brandt) 

Thalassema (Echiurus) sitkaensis Brandt, 1835, p. 62. 
Echiitrus silkaeyisis Spengel, 1912b, pp. 184-189. 

Diagnosis. — Corpus circiter tripollicarc oblongum, e subbrunneo 
olivaceum, obscurius punctatum et transversim striatum. Proboscis 
latiuscula, carnea, transversim purpureo striata, apice emarginata. 
Unguiculi anterioris corporis partis et spiculac posterioris lutea. 
(Brandt.) 

Differing from E. echiurus in having tw^o nephridia, in lacking a 
differentiated ridge of tissue along ventral side of proboscis, and in 
having skin papillae subequal rather than in rings of larger papillao 
alternating with narrow zones of smaller. 

Remarks. — This species constitutes one of the major mysteries in 
the systematics of the Ecliiuroidea. Ivlertens collected two specimens 
at Sitka, both of wliich he dissected. One of these specimens, his 
notes on the dissections, and a life sketch in color reached the St. 
Petersburg Museum and were used by Brandt. Subsequently all 
these became available to Spengel, as he details in his Echiurus paper 
(1912b). 

Spengel made a thorough examination of what remained of the 
internal anatomy and was able to satisfy himself that only two nephrid- 
ia were present, in the location of the anterior nephridia of E. echiurus. 
The proboscis was very adherent to the body, and it lacked the ridge 
of tissue on its concave under surface. As less important differences 
he lists: skin papillae subequal, in rather regular and very numerous 
rings (not rings of larger pa])illae alternating with zones of smaller); 
curvature of anal setae stronger than in echiurus; color, according to 
Mertcns's drawing, browiiish olive spotted and cross-stripad with 
darker, the proboscis flesh color with i)ur])le transverse stripes. 

The tougli, nondcciduous proboscis is characteristic of the Alaskan 
I'lchiurus I have examined, but all these have the ventral ridge 
I)resent, leaving as the princi])al characters of sitkaensis the two ne- 
phridia, absence of proboscis ridge, and the subequal papillae. 

Wilson (1900, p. 174) states that he examined Alaskan specimens 
of E. echiurus { = ala.skanu.'i) collected by J)r. W. R. Coo in 1899. 
"This species was found abundantly at numy different localities 
along the Alaskan coast south of the Peninsula and on adjacent 
ishuids, nearly always in rich black mud." I have listed 120 speci- 
mens from Alaska and British Columbia. None of these is sitkaensis. 

If there ii.v a sj)ecies sitkaensis it may nonnally live below low tide 
and only occasionally be carried shoreward during heavy storms. 



230 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

Type. — Formerly in St. Petersburg Museum; collected by H. 
Mertens. 

Type locality. — Sitka, Alaska. 

Genus THALASSEMA Lamarck 

Thalassema Lamarck, 1801, p. 328 (type, Lumbricxis thalassema ' Pallas, 1771, 
Spicilegia Zoologica, fasc. 10, p. 8, pi. 1, fig. 6). 

Diagnosis. — Echiuridae with a well-developed proboscis but \\dthout 
anal bristles and without specialized bands in the longitudinal layer 
of body muscles; inner oblique layer smooth, except sometimes for a 
short distance at anterior end of body ; anterior nephridia (gonothecae) 
one or two pairs, the internal ciliated funnel (nephrostome) without 
spiral lobes. ^ 

THALASSEMA STEINBECKI, new species 

Figure 11 

Diagnosis. — Small, slightly translucent, the proboscis as long as 
body, broad proximally, ribbonlike distally; nephridia, two pairs, the 
cUiated funnel with simple subcircular opening lacking any trace of 
sphal lips; interbasal muscle of setae well developed, strong, passing 
through loop of dorsoventral blood vessel; siphon beginning a short 
distance from vascular ring; precloacal intestinal coecum; intestinal 
mesenteries including conspicuous subfusiform fleshy masses; anal 
vesicles as long as contracted body, covered with numerous tiny 
ciliated funnels. Length of body 12 mm.; of proboscis, 12 mm. 

Description. — The skin is coarsely verrucose for a short distance 
back of proboscis and on terminal tliird of body; elsewhere the verrucae 



' Thalassema was first used in a generic sense by Lamarck. Although Pallas (Spicilegia, 1771) mentions « 
the name Thalassema as used by Joseph Gaertner, he names the animal Lumbricus thalassema, which appears 
to be the first valid binomial referring to the species generally known as Thalassema neptuni Gaertner. The 
latter is in effect a manuscript name. Neither Shipley (1899, p. 351) nor Wharton (1913, p. 265), who have 
offered revisions of Thalassema, gives any reference for the combination Thalassema neptuni. Quatrefages 
(1865, vol. 2, p. 595) cites "Thalassema Neptuni Gaertner, citfi par Pallas, Spicilegia Zoologica, fasc. 10, p. 8, 
pi. 1, fig. 6." Forbes, 1841, in his "History of British Starfishes and Other Animals of the Class Echino- 
dermata," gives a good account of the habits of "Gaertner's spoon-worm, Thalassema Neptuni Gaertner." 
The first item in his list of references is Lumbricus Thalassema Pallas. 

The derivation of the word seems to be thnlassos (sea) + ema (dart). 

8 Shipley (1899, p. 351) in his revision of Thalassema is in error in the statement that the nephridia have 
their internal openings gpirally twisted. Lankester (1881, p. 355) writes that they are semicircular and 
contrasts them with the spiral sort found in T. moebii, as figured in Greef's "Die Echiuren" (1879, pi. 8, fig. 69). 

Figure 11.— Thalassema steinhecki, new species: A, The type, X 14, showing arrangement of 
organs; the greater part of the intestine has been omitted and the foregut has been strongly 
bent to right in order to uncover the nephridia. B, A seta, X 20, from a specimen taken 
in 165 fathoms off San Francisquito Bay, Baja California; above, the hook, further en- 
larged. C, Ventral view of the type, X 4. {AV, Anal vesicle; B^-B*, dorsal, ring, neuro- 
intestinal, and ventral vessels; C, stomach; CG, ciliated furrow of intestine; CI, cloaca; 
G, gizzard; /, intestine; IC, intestinal coecum; Me, mesenterial bodies; MI, interbasal 
muscle of setae; N, nephridia; NC, nerve cord; 0, esophagus; P, pharynx; S, seta; Sil, 
anterior end of siphon.) 



ECHIUROID WORMS OF NORTH PACIFIC — FISHER 231 




FicuRK 11. — (See opposite pa^e for legend). 



232 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

are obviously smaller, and there is a suggestion of transverse align- 
ment. The appearance of the skin will depend a good deal on the 
state of contraction of body wall. The middle portion of body is 
slightly translucent, not transparent in places as in Listriolobus pelodes. 

The setae have a stout interbasal muscle connecting the inner ends, 
from which radiate numerous fan-shaped sheets of muscle which 
attach broadly to the body wall. A posterior muscle passes over the 
first nephridium and merges with the body wall between the first and 
second nephridia. Setae 1.8 mm. long with a small hook, the point 
of which is at right angles to the shaft. In the type and a number of 
other specimens the hook is worn off. 

Inner layer of muscles of body wall smooth except in front of 
anterior nephridia where there is differentiation into about a dozen 
bundles passing around the body. The middle layer is undifferen- 
tiated as in other species of the restricted genus Thalassema. 

The four nephridia are small in the type, which is probably a male. 
In the specimen from station 2828 they are enormously inflated and 
full of eggs. In the type the ciliated funnel is very simple, with 
smooth lips forming an incomplete circle. In the female the funnel is 
flattened and pear-shaped and somewhat distorted by pressure, but 
there is no indication of elongation into spirals. 

The deflated anal vesicles are as long as the body and have the 
appearance of being capable of inflation to a great size. 

The alimentary canal is moderately long but conspicuously shorter 
than that of Listriolobus pelodes. The contents are not always 
formed into pellets. The foregut consists of a pharynx-esophagus, 
midway along which is a sharp bend. Both the gizzard and stomach 
are unusually short. The former is marked by ring striations. At 
the posterior end of the stomach is the ring blood vessel (5^) marking 
the beginning of the intestine, on the lower side of which is the ciliated 
groove. At a distance back of the ring vessel equal to about length 
of gizzard and stomach the siphon begins and runs for about half 
length of intestine. The ciliated groove continues to the intestinal 
coecum in front of the cloaca. The pharynx has a conspicuous 
ventral mesentery, and all along the canal are numerous dorsal 
mesenterial strands. Posterior to the region of the siphon the mesen- 
teries inclose, or hold, conspicuous yellowish-white masses, sometimes 
subfusiform or in irregular sheets, which remind one of the suet found 
in mammals. A squeeze reveals several sorts of cells, some of which 
may be immature sperm. The cloaca is small and thin-walled. 

The vascular system consists of the dorsal vessel ending posteriorly 
in the ring vessel, from which the neurointestinal connective passes 
to the ventral vessel, forming a loop enclosing the interbasal muscle 
of setae. The ventral vessel ends posteriorly at the intestinal coecum. 
In the female from station 2828 the vessels are the same. 



ECIIIUKOID WORMS OF NORTH PACIFIC — FISHER 233 

TVk-— U.S.N.M. No. 20G00. 

Type locality. — Ei Mogote, near La Taz, Baja California, low tide, 
March 22, 1940, 1 specimen, Steinbeck and Ricketts, 
Specimens examined. — Eight as follows: 

Albatross station 2S2S, Gulf of California, 2-1° II' 30" N., 109° 55' W., 10 fath- 
oms, shells, 1 specimen. 
La Plata Island, Ecuador, 7-10 fathoms, rocky with nuUipores, 2 specimens, 

Allan Hancock Foundation. 
Thurloe Bay, Baja California, S-10 fathoms, rock with gorgonids, 1 specimen, 

Allan Hancock Foundation. 
Off San Francisquito Bay, Baja California, 1G5 fathoms, shale and gray mud, 1 

specimen, Allan Hancock Foi^ndation. 
Ensenada de San Francisco, Baja California, 2-6 fathoms, 1 specimen, .\llan 

Hancock Foundation. 
.\gua Verde Bay, Baja California, 10 fathoms, mud and coral, 1 specimen, Allan 

Hancock Foundation. 
Dewey Channel, San Eugene Point, Mexico, 21-24 fathoms, coralline, rock, I 

specimen, Allan Hancock Foundation. 

Distribution. — Baja California to Ecuador, low tide to 1G5 fathoms. 

Remarks. — Mr. Ricketts states that the type was associated with 
living Denfalivm in sandy mud a short distance below the surface. 
The specimen from station 2828 has the intestine filled mth small 
fragments of shells. 

As this species belongs in the restricted genus Thalassema, it 
naturally resembles Th. thalassema. The gizzard, stomach, and 
presiphonal intestine are definitely longer in the latter species and the 
anal vesicles smaller, although with such extensible structures it is 
difficult to make comparisons. It may be recorded that m alcoholic 
specimens there is no division of the body into three parts, wliich 
Lcigh-Sbarpe (1928, p. 501) reports as a characteristic of thalassema. 
The ecology of the two species is very different. Lankester (1881. 
J). 350) found thalassema on the south coast of Devonshire "in galleries 
excavated in tlie red sandstone (not limestone) which is exjiosed at 
spring tides. The galleries appear to be those formed by the Lamel- 
Hbranch Gastrochaena which the Thalassema appropriates." Leigh- 
Sharpe (1928, {). 499) reports tlu- species from borings in hmestono 
made by the mollusk Saxicava, Fl^^mouth Sound. 

Named for John Steinbeck, whose expedition to the Gulf of Cali- 
fornia collected (lie type. 

Genus LISTRIOLOBUS W. Fischer 

Lislriolobua Spenokl, 1012c, p. 310 {noitien nudum). — W. Fisciiku, 1920n, p. 210 
(no type). (Type, Listriolohux hahamcnsia Fischer.) 

Diagnosis. — DifTcring from Thalassema, sensu stricto, in having 
elongate, s|)irally (H)iled lips to neplirostome and 8 to 10 narrow merid- 
ional thickening.s of the middle, loiigitudinnl muscle layer. Differing 
from Ocheiostoma in having the inner oMiciue layer a smooth con- 



234 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

tinuous sheet between the longitudmal bands, not divided into 
separate fascicles. Nephridia 2 or 3 pairs; interbasal muscle of setae 
present. 

The character of the longitudinal muscles is not well marked in L. 
pelodes until the animal has reached a length of 15 to 20 mm. and 
after it is sexually mature. Apparently in L. sorbillans (Lampert) a 
similar condition exists (Wharton, 1913). Even so, the structure of 
the nephrostome will segregate these species from true Thalassema. 
L. riukiuensis (Sato, 1939, p. 359, figs. 10-13) is probably not a 
Listriolobus. There is no interbasal muscle to setae; the diagram of 
the blood vessels is not the Listriolobus pattern. 

Spengel (1912c, p. 316) established Listriolobus for Thalassema 
erythrogrammon of Sluiter (1883) and of Wilson (1900). Sluiter's 
species came from BHliton in the Java Sea whUe Wilson's was taken 
in the Bahamas, Spengel had Sluiter's animal and a duplicate, from 
Florida, of Wilson's species, which he characterized as "nearly re- 
lated." Unfortunately, as neither of these species had a valid name, 
Listriolobus was without a type and was technically a nomen nudum, 
Spengel did mention Thalassema mellita Conn as a species inquirenda, 
"which species one must include in the above genus I can not at 
present decide, nor even whether or not it is justifiable to include 
such a form as Th. mellita, which has bundles of longitudinal muscles 
that arise from thickenings of the continuous longitudinal muscle 
layer." A few lines farther on he says: "The animals described by 
Sluiter and Wilson as Thalassema should be put in the same genus 
with Th. mellita because of the nature of the sheaths surrounding the 
tunic muscles. I propose the generic name Listriolobus for the species 
of Sluiter and Wilson." 

Wilhelm Fischer (1926a, p. 110) discusses Listriolobus and names 
Wilson's and Sluiter's species Listriolobus bahamensis and L. billi- 
tonensis. So far as I know this is the first association of Listriolobus 
with a species from which a type can be chosen. Since Fischer did 
not do this I will so designate Listriolobus bahamensis Fischer, as 
being the species more likely to be available for study in the future. 
By implication Fischer includes Th. mellita Conn in Listriolobus, but 
I have made it the type of a new genus, Lissomyema. 

LISTRIOLOBUS PELODES. new species 

Figures 12, 13; Plate 21, Figures 1, 2, 4, 4a, 4b; Plate 22 

Description. — Largest specimens 40-60 mm. long and 12-25 mm. 
thick; proboscis capable of extension to slightly exceeding length of 
body, narrow, thin, translucent, so that nerve loop is visible. 

Body wall tanslucent, marked by eight narrow longitudinal muscular 
thickenings which appear light gray agamst the darker and much 
broader areas between them. There are a middorsal and midventral 



ECHIUROID WOKMS OF NORTH I'ACllTC — FISHER 235 




Figure U— Listriolobus pelodtj, new spoics: Small 8j)ccimcn, X IS, showing arrangement 
of organs. The greater part of the intestine has been removed. In this specimen the 
ncphridia arc very small. Lettering as in figure II. 



236 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

band and three lateral, equidistantly spaced. The bands, 1.5 to 
2 mm. broad, represent concentrations of longitudinal muscle fibers, 
which gradually thin out on the sides of the bands. The skin is 
beset. with small, unequal, subcircular and elliptical glandular thick- 
enings arranged in transverse close-set lines. At posterior end there 
is an area of greatly enlarged papillae. Coelomic surface of body 
wall smooth; innermost layer of slightly oblique muscle fibers very 
thin, uniform, not mterrupted by the longitudinal thickenings of 
the middle layer. 

Setae 2, close together, and close to the mouth. Sometimes two 
setae occupy one of the sheaths, with a single normal one in the 
other. There is an interbasal muscle uniting top of the sheaths. This 
passes through a loop in the dorsoventral blood vessel. 

Nephridia 4, variable in size. In the type the anterior pair is 
smaller than the posterior, but in another specimen the anterior 
pair is the larger and the four are about five times the length of those 
of type. They have the same terminal slender portion, which can 
undoubtedly be expanded. Both specimens are males. The anterior 
pair is situated posterior to setae about the length of the latter. The 
ciliated funnel has long coiled lips and is attached to the outer side 
of the base of nephridium by a short stalk. In a third specimen the 
nephridia are empty and reduced to filaments slightly expanded at 
base. In a full-grown female (Tomales Bay) the nephridia are 20 
mm. long, slender, and contain numerous eggs (June 7), 0.08 to 0.09 
mm. in diameter. 

Anal vesicles 2, variable in size, capable of great extension. Each 
is fastened to body wall about 5 mm. laterally from the anus by one 
or two mesenteries and ventrally by another pair close to nerve. 
Into these, which appear hollow, extends a short diverticulum of 
the vesicle. There are scattered, very tiny, ciliated funnels. The 
well-preserved female from Tomales Bay has vesicles that extend as 
far forward as the large posterior nephi'idia and lack the basal diver- 
ticulum found in the male. 

Alimentary canal. The pharynx extends to the sharp bend shown 
in the illustration, followed by the esophagus. The gizzard is short 
and the stomach relatively long, but there is bound to be variation in 
different specimens o^ving to the accidents of preservation. The 
intestine proper starts just back of the ring blood vessel, at the 
beginning of the ciliated groove, which soon becomes differentiated 
into the siphon. Even in the carefully hardened specimen from To- 
males Bay the intestinal wall is delicate and transparent. The 
length is difficult to measure on account of unequal contraction. The 
siphonal part is on the order of 100 mm. in length and the postsiphonal 
125 mm. (length of specimen, 40 mm.). Throughout its length the 
intestine is stuff ed with small unequal (1-2 mm.) ellipsoidal mud pel- 



ECIIIUHOID WORMS OF NORTH PACIFIC FISHER 237 




Figure 13. — Listriolobuj pdodes, new species: A sexually mature but small female, showing 
nephridia greatly swollen by contained eggs (not indicated), X 15. Lettering as in 
figure 11. 



238 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

lets. Just before the liind-gut passes into the small cloaca there is a 
spherical ventral coecum, to which extends the ventral blood vessel. 

The vascular system can be readily followed on plate 22. The 
neurointestinal connective forms a loop around the interbasal muscle 
of the setae. The two flaplike expansions of this vessel, shown in 
the figure, are not present in the Tomales Bay specimen. The dorsal 
vessel varies widely in diameter being sometimes greatly inflated 
(Tomales Bay), the inflation extending into the neurointestinal 
connective. 

Owing to the very thin, translucent body wall the nerves can some- 
times be seen under strong illumination such as sunlight. They pass 
directly around the body, from the ventral nerve cord, without visible 
branches. The translucent proboscis affords an opportunity to trace 
the proboscis loop throughout its entire course (pi. 21, fig. 2), This 
ganglionic continuation of the ventral cord is near the margin of the 
proboscis, to which numerous tiny nerves pass from slight ganglionic 
thickenings on the outer side of the cord. No nerves, under favorable 
conditions, could be detected on the mesial side of the cord. 

Color in hfe: "Proboscis yellow orange, deepest on edge; body dull 
grey-violet with greenish raised specks about 0.5 mm. in diameter, 
spaced quite regularly about 0.5 mm. apart (the papillae); 8 Ughter 
colored muscle bands" (large specimen from Tomales Bay). 

Small phase (figs. 12, 13; pi. 21, figs. 4, 4a-b). Small examples 
are fairly common on muddy bottom in moderate depths off southern 
CaHfornia and in Newport Bay. These preserve badly unless special 
care is exercised. They are usually strongly contracted into sub- 
spherical form and vary greatly in appearance. Sometimes the body 
wall is rather uniformly transparent but more often the posterior 
portion is opaque. This region may be smooth or thrown into eight 
meridional swellings which occasionally extend the whole length, 
giving the appearance of a tiny melon. These swellings are caused 
by the contraction of the eight muscle bands and are found in strongly 
contracted specimens 7 mm. long. But specimens in which the body 
wall is stretched and transparent do not show indications of differ- 
entiated muscle bands until much later — at a length of about 15 to 
20 mm. In the very small sizes the muscles are Ukely to show first 
posteriorly and ventrally. A specimen 20 mm. long from 55 fathoms 
off Santa Cruz Island has the 8 bands fully developed. Even in large 
specimens (40 mm.) the bands are not always equally conspicuous. 

I have found a specimen 7 mm. long sexually mature. Normally 
these small examples are transparent in the midregion so that the 
nerve cord, intestinal pellets, and egg-laden nephridia can be plainly 
seen as yellowish bodies in life. The intestine is characteristically 
thin-walled, transparent, and highly convoluted and taxes the capacity 
of the body cavity. Its walls are greatly distended with mud peUets 



ECHIUROID WORMS OF NOHTH PACIFIC — FISHER 239 

about 1 mm. long. A specimen from Los Frailes, Baja California? 
has the intestine distended with sand not in pellets. 

The specimen shown in plate 21 , figure 4, was 12 mm. long and 9 mm. 
liiick when alive, contracting to about 10 by 10 mm. when killed. 
The transparent areas of body wall are characteristic and are dotted 
with gi'ayish or wliitish papillae. In life the general tone is greenish 
or olive, with gi'ay papillae spots, the proboscis bordered with yellow. 
In the clear area the lateral nerves can be seen as they leave the ven- 
tral nerve cord and in favorable specimens the nerve loop can be 
traced around the entire margin of proboscis. 

Dr. Olga Hartman describes small specimens from off southern 
California as being rich, dark, satiny green in life, wliile those from 
Newport Bay Professor MacGinitie found to be decidedly greenish. 

Text figures 13 and 14 indicate how widely the neplu-idia vary in 
size. In the swollen state the wall is perfectly transparent and 
excessively thin and the proximal parts of the four vesicles adhere to 
each other, the distal part lying at random, crowded amid the close 
coils of the mud-filled intestine. The eggs are suspended in a thin 
gel, spaced 1 or 2 diameters apart. They vary from 0.09 to 0.1 mm. 
in diameter and are gi-avish in color, surrounded bj^ a clear zone 
(about 0.009 mm. thick). The clearly visible nucleus is 0.045 mm. in 
diameter. 

In text figures 12 and 1 'A the vascular system is shown in a contracted 
state. It does not (lifFcr in essentials from that of the fully adult 
(pl. 22). 

7;v/>^.— U.S.N. M. No. 20008. 

Tyj)e locality. — Monterey Bay, Calif., moderate depth, fine sand; 
from stomach of flounders. 

Specimens examined. — As follows: 

Monterey Bay, 10 large and 4 small specimens. 

Tomalcs Bay, Marin County, Calif., low tide, soft black sandy mud, about 6 

inches below surface, 1 large female, June 7, 1941 (University of California). 
Newport Bay, Calif., 7 to 20 fathoms, numerous specimens, 4 to 1 5 mm. ; collected 

by Prof G. E. MacCinitio. 

The Allan Hancock Foundation tendered for examination an impor- 
tant collection from Baja California and off southern California, com- 
j)rising the small phase and some of intermediate size (20 to 40 mm.): 

JiAJA Camkornia 

Lfis p'raih-s, 5 1.') fathoms, sand aixl nlnac, 1 specimen. 

I'ppcr end of Culf of ("aiifornia, 21 fathoms, brown mud, 6 specimens. 

Camkciuma 

Off Newport Beach, ."iO fathoms, mud, 7 specimens. 

Off Laguna Beach, 2.'> .'>7 fathoms, sandy mud, 2.') specimens. 



240 PROCEEDINGS OF THE NATIONAL MUSEUM vol. sa 

Off Bluff Cove (8 lots) 25-100 fathoms, mud, fine sand, 32 specimens. 

Off Redondo Beach (15 lots), 10-120 fathoms, mud, fine sand, coarse sand and 

mud, 44 specimens. 
Off Point Vicente Lighthouse, 17-40 fathoms, coarse sand and mud, 4 specimens. 
Off Portuguese Point, 16-20 fathoms, gray sand and seaweed, 2 specimens. 
Cortes Bank, 60 fathoms, sand, broken shell, 3 specimens. 
Off Point Mugu, 26-30 fathoms, mud, 2 specimens. 
Santa Catalina Island, 50-51 fathoms, mud, 3 specimens. 
Santa Cruz Island (5 lots), 31-138 fathoms, mud, sand, 15 specimens. 
Santa Rosa Island, 28-45 fathoms, 5 specimens. 
San Miguel Island, 35 fathoms, mud, 20 specimens. 

Remarks. — The type and largest specimens were recovered from 
the stomach of flomiders from moderate depths of Monterey Bay. 
In nearly all the examples the intestme is disintegrated and the body 
cavity filled with pellets of fine sand. This is true also of specimens 
from southern California, which were dredged and placed almost at 
once into alcohol. 

After the drawings and description were completed I received from 
Prof. S. F. Light a carefully hardened specimen from Tomales Bay, 
collected at low tide, in mud frequented by the clams Schizoihaerus 
nuiialli and Macoma secta. This specimen (June 7) was apparently 
laying eggs as the nephridia are partly emptied. It is especially valu- 
able as it gives the characters of the adult female, is much better pre- 
served than the Monterey specimens, and affords opportunity for 
recording the life colors of a large example. 

In the type specimen, near the anterior left nephridium is a light- 
colored lobed mass (pi. 22, x) adherent to body wall. It is possibly a 
parasite. 

Genus OCHETOSTOMA Leuckart and Riippell 

Ochetostoma Leuckart and RtJpPELL, 1828, pp. 7-8 (type, 0. erythrogrammon 
Leuckart and Riippell). — Spengel, 1912c, p 316. 

Diagnosis. — Greater part of the thickness of the longitudinal muscle 
layer segregated into separate longitudinal bands. The intervals 
between these bands is crossed by very niunerous separate small 
muscle bundles of the inner oblique layer, which remains for the most 
part continuous and unbroken over the surface of the longitudinal 
bands. Anterior nephridia, 1 to 4 pairs, the coelomic aperture having 
spirally coiled lips; with or without coecum at end of intestine; inter- 
basal muscle of setae present or absent; two ventral anterior hooked 
setae; no anal setae. 

Remarks. — The list of species that follows is mostly derived from 
literature and is in nowise intended to be a revision, for which speci- 
mens and new dissections will be absolutely essential. Most of the 
species were described as Thalassema. Th. exilii (Fritz Miiller) 
Lampert, which has 8 to 10 muscle bands and two pairs of nephridia, 



ECHIUROID WORMS OF NORTH PACIFIC — FISHER 241 

with nophrostomc lips merely folded and crinkled, but not elongated, 
is possibly a Lissomyema. 

a^. Five pairs of nephridia, 3 rather poorly developed, in front of setae; 19 muscle 

bands -- O. homelli (Prashad) 

Five or four pairs of nephridia, 2 in front of setae; 10 or 11 muscle bands. 

O. bombayensis (Prashad and Awati) 
a'. Four pairs of nephridia, posterior to setae. 

10 muscle bands O. decameron Lanchester 

20 muscle bands O. kempi (Prashad) 

a}. Three pairs of nephridia, the first pair opening in front of the ventral setae 
the second and third pairs posterior to setae. 

14 muscle bands O. erythrogrammon LcuoJcart and Ruppcll* 

15 or 16 muscle bands O. stuhlmanni (Fischer)' 

15 to 17 muscle bands O. leptodermon (Fischer)' 

16 to 18 muscle bands O. caudex (Lampert)' 

17 or 18 muscle bands O. kokotoniense (Fischer)' 

17 or 18 muscle bands O. grifflni Wharton 

a*. Two pairs of nephridia. 

7 or 8 muscle bands; papillae white, scattered uniformly all over body; 
color white in alcohol; proboscis one-third to one-half length of 
body; anal vesicles broad and saclike O. formolosum (Lampert) 

8 muscle bands. 

With incipient gills on edge of proboscis at base; broad smooth zone 

around anus arkati (Prashad) 

No branchial fringe to proboscis; no smooth zone at posterior end 
of body; color in life greenish, in alcohol grayish flesh color; 
proboscis one-third to length of body. 

octomyotum, new species 
10 or 11 muscle bands O. hupferi (Fischer) 

13 muscle bands; proboscis longer than body; anal trees one-fourth length 
of body O. pellucidum (Fischer) 

14 muscle bands; anal trees more than one-half body kiigth. 

O. manjuyodense Ikeda 

17 to 19 muscle bands; proboscis shorter than body; color dark green 

with violet longitudinal stripes and with white papillae scattered all 

over the body; anal vesicles long, brown, pointed anteriorly, bearing 

short branched outgrowths O. baronii (Greef) 

16 or 17 muscle bands; proboscis deciduous, can equal length of body; 
setae with interbasal muscle; anal vesicles without branched 
outgrowths... _.0. edaz, new species 

OCHETOSTOMA OCTOMYOTUM. new Bpcclcs 
Plati: 21, Fun;uK 3; Platks 23, 24 

ViagnoHu. — Skin transluoent, with ei^ht more opatiiic" lonpitudinnl 
strifx'S marking the inuselc bands, and closely sti|>ple(l everywiiere 
witii very small, une<|ual, often elliptical glands wliieh increase in 
size very nuirkedly over the j)osterior region (where tiie niusele bands 
fuse into a continuous sheet); i)roboseis thick and fleshy in preserved 

• Sato, 1930, p. Vil considers these flvc, bIopk with Thnlntitma palmie lkr>Iii, Urjl, to I'O nil ono sprcii'M. fdr 
which trythronTammon U the oldest name. 



242 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

specimens, varying from less than one-third to the full length of body. 
Four nephridia, the nephrostome with very long coiled lips. Siphon 
begins a short distance behind the ring blood vessel (marking the 
boundary between foregut and intestine) ; anal vesicles long, with 
pointed apex and with tiny ciliated funnels scattered over surface; 
setae without interbasal muscle. Length of full-grown specimen 110 
mm. ; proboscis 30 mm. ; diameter 20 mm. A specimen collected at 
Cabrillo Beach, San Pedro, Cahf., is 95 mm. long; proboscis 93 mm. 

Description. — Longitudinal muscle bands 8, well developed, broader 
than the intervals between, rather iridescent and situated as a mid- 
dorsal, midventral, and (twice) three laterals. These fuse into a 
continuous sheath on the posterior part of body, equal to about one- 
third body length of expanded specimens and less of contracted ones. 
Anteriorly they remain separated to the base of proboscis. The 
innermost or oblique muscles are well developed, and between the 
longitudinal muscle bands they form a consecutive series of oblique 
partitions alternating with narrow compartments. At the bottom 
of these compartments is a thin layer of the longitudinal middle sheet. 
In the posterior region the oblique fascicles become less and less 
distinct as the anal area is approached. In reality the uninterrupted 
layer of longitudinal muscles is covered by a continuous inner layer 
of oblique muscles differentiated into thicker and thinner portions. 
Thus, forward from the anus the gradual differentiation of the separate 
oblique strands can be easily seen. In severely contracted specimens 
this posterior region does not show to advantage. 

Anterior nephridia, 2 pairs, the ducts piercing the midventral 
muscle about midway between its outer margin and the nerve cord. 
The anterior pair is situated a short distance behind the setae (rather 
less than length of seta). The interval between first and second is a 
little more than twdce the distance. They open internally by ciliated 
funnels having very long, coiled, extensible lips. 

Anal vesicles 2, unbranched, capable of great distension. They open 
on the ventral surface of cloaca. Tiny scattered ciliated funnels 
may be seen on the surface. 

Alimentary canal. There are three general regions: (1) An 
anterior division or foregut, in which the longitudinal musculature lies 
outside the circular musculature; (2) an intestine proper, in which this 
order is reversed, characterized by the presence of the ciliated groove 
and along a part of its course by the collateral intestine or siphon; 
and (3) a short rectum, or cloaca, posterior to the coecum. 

The foregut consists of the pharynx, esophagus, gizzard, and stom- 
ach, and the junction with intestine coincides \vith the position of the 
ring blood vessel. The pharynx has a tough wall, rather iridescent 
with wavy longitudinal lines; the esophagus is thin-walled and con- 
voluted, while the gizzard and stomach, both brief, can be differen- 



ECHIUROID WORMS OF NORTH PACIFIC — FISHER 243 

tinted by texture. The gizzard is nmri-ced by eircular lines, the stomach 
by longitudinal representing the longitudinal folds of the lining. 
Complicated dorsal and ventral mesenteries attach the pharyiLX and 
esophagus to body wall. 

The hitestine is, as usual, highly convoluted but when partly un- 
raveled is seen to have a principal posterior loop, and then an anterior. 
The siphon, or collateral intestine, begins a short distance behind the 
ring blood vessel, its course being shown in plate 23, figure 1, where 
the dotted line indicates the ciliated groove, which terminates at the 
coecum. The walls of the cloaca have thick longitudinal folds. The 
anal vesicles empty into it by veiy small pores. 

Blood-vascul.\r system. The dorsal vessel, intestinal ring vessel, 
doi-soventral comiectives, and ventral vessel are shown in plate 24. 
The ventral vessel, which anteriorly follows a part of the free edge of 
the ventral mesentery, terminates posteriorly on the wall of the 
coecum. 

Color in life, greenish; in alcohol grayish flesh color with eight dark 
grayish stripes. 

fyi^f.— U.S.N. M. No. 20607. 

Type locality. — Newport Bay, Orange County, Calif., January- 
Februarj^ 1930, G. E. MacGinitie, 11 specimens. 

Other material examined. — Cabrillo Beach (San Pedro), Calif., 1 
specimen, W. R. Lloyd, Cabrillo Beach Marine Museum. 

Remarks. — Newport Bay, where Professor MacGinitie collected 
tho tyi)e series, is a rather small tidal inlet south of Long Beach. 
The worms were taken from a sandy bar, exposed at low tide, where 
they inhabited U-shaped burrows, the mouths of which were 12 to 14 
inches apart and the bottom 10 to 12 inches below the surface. A 
mutilated specimen was dredged in 25 fathoms. 

This sand bar was later removed in order to deepen the harbor, and 
tiie species was apparently wiped out at that locality as it has not since 
Incn taken in spite of nmch collecting. 

Tlie food consists of very fine detritus, which in (he intestine is 
fornu'd into ellipsoidal pellets 2 to 2.5 mm. long by 0.75 mm. thick. 

The only species with which 0. octomyotum might be confused is 0. 
formolosvm (Lampert) from the Pliilippincs and Shanghai. This is a 
small form averaging 30 mm. in length, |)r()boscis 8 mm., and diameter 
10 mm. The color is white in alcoholic sjx'cimens, the skin very 
thin, with white papillae scattered uniformly all over the body. The 
anal vesicles are described as broad saclike organs. The species has 
seven or eight muscle jjundles, two pairs of nephridia with spirally 
coiled openings, and a .spherical diverticulum on the rectum — charac- 
tcri-.tics of odorinjotum, except the nunil)(r of nmsclcs which is con- 
stantly eight in the California si)ecies. 



244 



PROCEEDINGS OF THE NATIONAL MUSEUM vol. 




Figure 14. — Ochetostoma edax, new species: A, Dissection of anterior portion of body to 
show relations of nephridia, setae, vascular system, and anterior part of alimentary canal, 
X 6. B, Ventral view of specimen from Puerto Refugio, X 2. (TV, nephridium similar 
to the four of type, a male; N^, female nephridia from a smaller, Puerto Refugio, speci- 
men. Other lettering as on plate 24.) 



ECHIUROID WORMS OF NORTH PACIFIC — FISHER 245 

The specimen from the Cabrillo Marine Museum was taken in sand 
and is notable for the 93-mm. proboscis, whicli is IG mm. broad near 
tne base. The nerve loop, following the margin, is plainly visible. 

OCHETOSTOMA EDAX, new species 

Figure 14 

Diagnosis. — Two pairs of nephridia situated posterior to setae, the 
nephrostome with spiral lips; setae with interbasal muscle passing 
through a small loop of the neurointestinal blood vessel; longitudinal 
muscle bands 16 or 17, with very narrow interspaces; rather long pre- 
siphonal intestine; precloacal coecum; two unbranched anal vesicles 
capable of great distension. Body wall translucent in middle region, 
striped with narrow dark zones; skin papillae numerous, sometimes 
whitish, in not well-defined transverse lines, and usually larger on 
posterior third or fourth of body, but not hard to the touch; proboscis 
deciduous, fleshy, from one-fourth to body length. Length of body 
25 to 50 mm. 

Description. — The IG or 17 muscle bands merge into a continuous 
sheet in ttie posterior fourth of body. The intervals between the 
bands are narrower than in octomyotum and transverse oblique fas- 
cicles of the inner layer are decidedly weaker. The ventral muscle 
band under the nerve cord is broader than the others. Sixteen is 
probably the normal number of bundles, the extra band arising from 
the incomplete splitting of one of the lateral or dorsal bundles. 

Nephridia 2 pairs, opening at outer margin of midventral muscle 
and varying greatly in size. In a male the nephridia are very large, 
the posterior reaching to posterior end of (contracted) specimen, 
while in a smaller female they have the relative size of the three 
smaller nephridia of figure 14. The ciliated funnel on the anterior 
face of uephridium has long coiled lips. The two anal vesicles, with- 
out branches, are covered with ciliated funnels and are capable of 
being distended to a large size. They are as long as the body in the 
largest specimen, which is, however, strongly contracted. 

Ali.ment.\ry canal. Forcgut relatively short, ending with the 
ring blood vessel (IP). Oidy two parts are clearly difi'erentinted, the 
phar}-nx-esophagus and stomach, with longitudiiud folds. The 
gizzard of 0. octomyotum appears to be lacking. The intestine is 
long, highly convoluted, witii a very much longer presiphonal segment 
than in octomyotum, being lojiger than the foregut. The wall is thin 
and in the type is greatly distended with coarse sand; in another 
specimen, with sizable fragments of shells and miscellaneous hard 
deV>ris. In the type the presiphonal intestijie measmes roughly 
30 mm., the siphonal part 120 mm. and postsiphonal 90 mm. The 



246 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

latter has a ciliated groove terminating at the coecum just in front 
of the cloaca. The walls of the coecum are much thinner than in 
octomyotum and lack heavy longitudinal folds. 

Blood-vascular system (fig. 14, B^-B^). This is on the same 
plan as in octomyotum, but in two specimens the left dorsoventral 
connective from the ring vessel was much longer than the right. 
Passing through the small loop where these connectives join the 
ventral vessel is an interbasal muscle of the setae, not present in 
octomyotum. The ventral vessel ends posteriorly in the coecum 
without branching. 

Color. "Specimens were elongate to grape-shaped, smooth and 
thin-skinned, greenish, with obvious and comparatively large spoon- 
shaped proboscis" (Steinbeck and Ricketts). 

T2/2?e.— U.S.N.M. No. 20606. 

Type locality. — Gulf of California: Pichalingue Bay, near La Paz, 
Baja California, February 1920, Luis G. Rubio. 

Specimens examined. As follows: 

Coronado Island, Gulf of California, March 27, 1940, Steinbeck and Ricketts, 
3 specimens, under and among slightly subtidal rocks on white sand. 

Point Lobos, Espiritu Santo Island, Baja California, March 3, 1940, Steinbeck 
and Ricketts, 1 specimen, under boulders of tide flats. 

Puerto Refugio, Angel de la Guarda Island, April 2, 1940, Steinbeck and Ricketts, 
10 specimens; under boulders on beach. All these are smaller than the type. 

Same locality, Allan Hancock Foundation, 1 specimen. 

Remarks. — The type is without proboscis. The sketch of the en- 
tire animal is from an example taken at Puerto Refugio. As some of 
the specimens lack a proboscis and others have only a small one it is 
probable that the organ is soon regenerated. The wide difference in 
size seems hardly to be due to accidents of fixation. 

The great disparity in size of the nephridia in the two specimens 
dissected shows that size is of no particular value as a character, 
depending as it does on the amount of conta'ined material. There is 
a wide difference also in the size of the anal vesicles in the two specimens. 

0. edax feeds upon the sand or coarser material in which it lives. 
This is not molded into definite fecal pellets. The specimen from 
Puerto Refugio had eaten very coarse material, which formed irregular 
masses in the intestine. Among the miscellaneous material could be 
recognized fragments of pelecypod and gastropod shells and small 
whole gastropod shells; serpulid tubes, calcareous bryozoans, barnacle 
shells; chelae of small crab; fragments of crab carapace, sea-urchin 
spines, and brown algae; many straight sUiceous sponge spicules and 
fragments of volcanic rock. Some of the sponge spicules were in 
bundles. Others had perforated the intestinal wall and were lying 
in the coelomic cavity. A few were in the anal vesicles. 



ECHIUROID WORMS OF NORTH PACIFIC — FISHER 247 

In the list of species, edax is next to bnronii, wliich has 17 to 19 
nuiscle banils and anal glands which are described as having short 
branching outgrowths. 

In July 191S I collected at English Harbor, Antigua, B. W. I., a 
small Ochetostoma that has 19 muscle bands, 2 pairs of nepliridia, 
a strong interbasal muscle passing tluough a loop of the dorsoventral 
blood vessel. The anal vesicles are long, reaching to the anterior end 
of the contracted specimen and are covered with ciliated cups, but 
there is nothing that can be interpreted as branched outgrowths. The 
proboscis is much contracted, fleshy, and very adherent. The 
posterior papillae are wliite, conspicuous, crowded, so that they are in 
contact, and are hard to the touch. In a section of the body wall they 
resemble calcareous nodules. This posterior part of the body is 
perfectly opaque while the rest is slightly translucent. This species is 
perfectly distinct from edax, and it is noted here because outwardly it 
would pass for 0. baronii. The specimen is a mature female with the 
four nepliridia enlarged and full of eggs. Length of body, much 
contracted, 20 mm.; thickness, 9 mm.; proboscis, 8 mm. 

Genus ARHYNCHITE Sato 

Arhynchite Sato, 1937, p. 142. (Type, Thalassema arhynchite Ikeda.) 

Diagnosis. — Differing from Thalassema in absence of a proboscis; 
nephridia 2, with spiral lobes to nephrostome; no intestinal coecum; 
no ring vessel around foregut at end of dorsal vessel, the connection 
between dorsal vessel and neurointestinal connective being indirect 
as in Bonellia; muscles of body wall smooth with no concentration of 
fibers in either the middle, longitudinal layer or in the inner, oblique 
layer. 

ABHYNCMITE INAMOENUS. new spcciea 

Platk 25 

Diagnosis. — Differing from A. arhynchite (Ikeda) in having a 
relatively longer neurointestinal blood vessel, which does not embrace 
the interbasal muscle. Intestine vciy long, especially presiphonal 
segment; dorsal blood ves.^el slender, not connecting directly with 
neurointestinal vessel, which is relatively long and divided into two 
before joining ventral vessel; size medium; skin with low papillae. 

Description. — Length of type, much contracted, 70 mm.; diameter 
25 mm. Skin roughened by low, close-set, pustulate verrucae largest 
and most irregular at QWiX^ of body. Setae 2, close to the much 
contracted anterior end; 11 mm. long, the inner ends united b}^ an 
interbasal muscle which docs not pass through loop of neurointestinal 
vessel, 

Coelomic surface of body widl perfectly smooth and with satiny 
sheen, the muscles continuous, without trace of difTerentiation into 
670329 — 1<; :; 



248 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 9« 

longitudinal or oblique bands. Where the seta muscles join body waU 
there are a few transverse thickenings of inner layer. 

Nephridia 2, inserted close to nerve and posterior to seta about 
the length of latter. In the only female dissected these nephridia 
are four-fifths length of body and contain numerous eggs. In the 
male they are about one-fifth length of body and contain sperm. 
The internal opening (nephrostome) is on a short peduncle near base, 
with an irregular small lip lacking any trace of spiral structure. 

Anal vesicles 2, simple, thin-walled, opening into small cloaca hav- 
ing longitudinally pKcate walls. Minute ciliated funnels are scattered 
over the surface. 

Alimentaiy canal excessively long (600 mm.) with very numerous 
coils attached to body wall by a multitude of very delicate frenula 
in which is entangled coagulum containing numerous eggs and brown 
bodies of unknown nature. The pharynx-esophagus is thin-walled. 
Plate 25, figure 3, shows the anterior complex more or less in situ 
and figure 4 with the interbasal setae muscle cut and the pharynx- 
esophagus pulled to right. The gizzard is about 6 mm. long, but the 
length of the stomach cannot be determined on account of condition 
of material. The interval between gizzard and beginning of siphon 
is the astonishing distance of 170 mm. The siphon accompanies the 
following 240 mm. of intestine, while the terminal, postsiphonal por- 
tion is 190 mm. There is no intestinal coecum in front of the cloaca. 

Vascular system. The dorsal vessel can be traced posteriorly 
nearly to the point where the long neurointestinal connective (B^) is 
attached to the lower side of the alimentary canal. The relation is 
similar to that of Bonellia. Anteriorly the neurointestinal connec- 
tion divides into two branches before joining the ventral vessel. 
There is therefore no enlarged "heart" at the posterior end of dorsal 
vessel, nor a ring vessel embracing the gut at that point. 

T^/pe.— U.S.N.M. No. 20615. 

Type locality. — Monterey Bay, Calif., 35-40 fathoms, mud, De- 
cember 3, 1931, 3 specimens. 

Other material examined. — Off southern California (13K miles south 
of Seal Beach), 215-225 fathoms, green mud, Allan Hancock Foun- 
dation, 1 specimen. 

Remarks. — The specimens are in a poor state of preservation, the 
alimentary canal being soft. 

The outstanding features of inamoenus are the two nephridia, the 
internal aperture of which does not have spirally twisted lips; the ex- 
traordinarily long presiphonal gut, and the very long neurointestinal 
connective not embracing the interbasal muscle of setae. The vas- 
cular system lacks a definite intestinal ring vessel — probably a generic 
feature. A. inamoenus differs from the only other recognized species, 
arhynchite of northern Japan, in respect to the neurointestinal vessel 



ECHIUROID WORMS OF NORTH PACIFIC — FISHER 249 

already mentionod, and in the probably much longer presiphonal gut. 
Details of the gut are lacking in Sato's figure and des('rii)tion. 

Family BONELLIIDAE Baird 

Bonellidae Baird (name only), 18G8, p. 111. (Includes Thalassema and Bonellia.) 

Diagnosis. — Dimorphic echiuroids. The male is degenerate, pla- 
narianlike, with ciliated ectoderm, generally one, exceptionally two, 
nephritlia '° serving as sperm receptacles, and a vestigial alimentary 
canal; it lives semiparasitically or parasitically on or in the female," 
and is sometimes absent from female; females resembling Thalassema 
but with bifid proboscis in some genera; two, four, or exceptionally 
many anterior ventral setae are sometimes present; no anal setae; 
anal vesicles with many branches ending in ciliated cups; one to three 
nephridia. 

KEY TO GENERA OF BONELLIIDAE 

a'. With an elongate proboscis bifid at the end. 

6'. With ventral setae " or hooks a short distance behind mouth, 
c*. Regularly one nephridium or egg receptacle (either right or left). 

d'. Coelomic aperture of nephridium (i. e., the nephrostome) situated 
near base of the organ, usually at end of a short lateral tube. 

Bonellia Rolando 
<P. Large nephrostome at extreme distal end of nephridium and not 

facing laterally Bonelliopsis, new genus (p. 252) 

c'. Regularly 2 nephridia, having the small nephrostome laterally near 
distal end; a small blind tube opening between nephridiopores serving 
as a permanent androecium for completely parasitic male; gonad of 
female situated on frenula radiating from cloaca; anal vesicles in form 
of tubules opening independently into cloaca (see also Acanthoha- 
mingia) Pseudobonellia Johnston and Ticgs '* 



'• Males of Pteudobontllia have two ncphri'lin; other penera one only. 

" The male lives In the foreput. In nephridiuni, on proximal portion of proboscis, in ponital proovc (/lean- 
thohamingia) . anrl In a si>cciallzed blind tube or androecium opening between the two nephridiopores {Pteu- 
dobontllia ) . 

'« In limellia miyniimai Ikeda (lOW, p. 73; 1907, p. 2, pi. 1, rtps. 1, 2; pi. 2, flps. 5-17) there are numerous 
(29) very small setae. Ptendobonellia has two to four sctiic; Archiboneltia has four and other pcnora have two 
Mtac, but sometimes o com piemen t/il smaller seta Is In process of prowth and liitcr replaces the fimrtlonal ono. 

'« John.ston and Tleps, 1910, pp. 21.V229, pis. »-ll. This is a very remarkable penus set ai)art from all other 
boncllilds in havlnp a small blind tube, projectinp into coelom and openinp on the ventral body wall between 
the two nephridloiwrcs by a narrow ainal whose walls contain stronp sphincter fibers. In this lives per- 
manently one very depenernto male. Its posterior end prown fast by onlarped ectodermal colls to the much 
smaller epithelial cells of the tubule. The male lacks setae and has two sperm receptacles, dlfTerinp from all 
other known males (which have only one). The ovary Is entirely dilTerent from that of other penera. "The 
mesenteric strands of mu.sculnr tl.ssue which nmlntain the iK)sterlor portion of the rectum in position arc very 
well dcvclopwl and form the basis of the ovary whilst from the ix-ritoneum llnlnp them the ova are developed" 
(p. 221). "The anal pinnds or pf»sterior nephridia are ropre,scnt/Ml by two small, tuft-like masses slliiate<l one 
on each side of the posterior end of Intestine. Each consists of n mass of very delicate, simple, cylindrical 
tubosopenlnpseparately Into the rectum, whose walls In thisreplon are thickened. The tubules arc approxl- 
mat/'ly clrailar in section with an Irrepular lumen. They consist of a sinple layer of epithelial cells. Near Its 
free end each tubule t)«comes narrowed before o[K>ninR Into the ooclom by a sllphtly dilated funnel fringed 
with long cilia" (p. 220). 

This species was collected In the Capricorn Group, Oreat Barrier Uoef, due cast from Kcppol Bay, Qaoens- 
land. Subsequently Monro (1931, p. 33) reported the species from Low Isloa. 



250 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

c^. Three nephridia, an unpaired between a pair; ovary along nerve cord; 
anal vesicles: a fascicle of tubules at end of a collecting tube. 

Archibonellia Fischer " 
6^. No setae present. 

c*. Nephrostome situated at end of a short lateral tube near blind distal end 

of nephridium Parabonellia Onoda 

c^. Distal end of nephridium expanded into plicated rim of large nephro- 
stome Eubonellia, new genus (p. 255) 

a^. Proboscis when present similar to that of Thalassema and not bifid at extremity. 
6'. No sharply marked groove between nephridiopore and mouth. 

c*. Female with 2 well-developed ventral setae; anal vesicles 2, elongate, 
dendritic; nephrostome near base of the single nephridium; male un- 
known Protobonellia Ikeda 

c^. Female with 2 ventral setae; nephridia 2, large, with basal 2-lipped 
nephrostome; anal glands wide sacs provided with a very large number 
of slender excretory tubules with apical funnel; male unknown. 

Maxmiilleria Bock 
c^. No ventral setae in female. 

rfi. Proboscis deciduous, Thalassema-like; 2 external papillae marking 
nephridiopores; nephridia 2 or 1 with basal nephrostome; anal 
vesicles in 2 thick clusters of tubules opening into a common duct; 

male with ventral hooks Hamingia Danielssen and Koren 

c*. In place of proboscis a short truncate snout; a proboscis possibly normally 
present; 1 nephridium with basal nephrostome; duct of nephridium 
passing under nerve cord and opening in median line into a funnel- 
shaped depression of skin; anal vesicles; numerous dendritic masses 
arising from a basal bladder on each side of large muscular cloaca; 
terminal portion of hind-gut, in front of cloaca, greatly enlarged; male 

unknown Nellobia, new genus (p. 257) 

^2. A narrow, or expanded, slit extending forward from nephridiopore, in 2 
species containing 8-10 tiny, integumentary spines; anal vesicles not in 
form of 2 dendritic structures or 2 clusters of tubules, but in form of 
independent branched tubes or an asymmetrical cluster; 1 or 2 nephridia 
with nephrostome near base; males with or without hooks. 

Acanthoharaingia Ikeda (p. 260) 



n Wilhelm Fischer, 1919, p. 283, figs. 5, 6; 1926b, p. 207, pi. 2, figs. 1-7, text figs. 1, 2. Archibonellia michael- 
soni, the type, from a coral reef, Rottnest Island, Fremantle, Australia, is only 12 mm. long; it has a large 
median between and above two small nephridia (no nephrostomes were found). The proboscis has two 
terminal lappets instead of divisions, and the alimentary canal is short, scarcely over twice length of body. 
A. mj6bergi (1926b, p. 208, fig. 1. northwest coast of Australia), however, has a very small unpaired nephridium 
(with basal nephrostome) between a pair of very large "uteri" filled with eggs and with terminal nephro- 
stome. The alimentary canal is of normal length for a bonelliid, and the proboscis is normally cleft at the 
tip. Fischer, in a quandry what to do with this species, places it in Pseudobonellia on the basis of the large 
paired nephridia, with terminal nephrostomes, and the normal alimentary canal and proboscis. Even though 
the androecium of Pseudobonellia may well have originated in an unpaired nephridium, the fact remains 
that in Pseudobonellia the androecium is highly specialized and is no longer a nephridium, while the ovary is 
not found along the nerve cord, as in Archibonellia, and the tubes of the anal vesicles open separately into the 
cloaca, not into a common duct. The two species of ^rcftiftoneHia may not be congeneric, but the aberrant 
species is certainly widely different from Pseudobonellia. 

Figure IS. — Bonellia viridis: Dissection (X 5) of anterior part of specimen from Naples 
to show particularly the position of nephrostome, CF, at base of nephridium, N, which was 
55 mm. long while the animal was only 45 mm. Note the long segment of gut between 
gizzard, G, and beginning of siphon, Sil. The anterior end of gonad, Go, Is shown. {B ^, 
B^, B*, dorsal, neurointestinal, and ventral blood vessels; C, stomach; G, gizzard; NC, 
nerve cord; 0, esophagus; P, pharynx.) 



ECHIUHOID WORMS OF NORTH PACIFIC— FISHER 251 

I 




Figure IS.— (See opposite page for legend}. 



252 



PROCEEDINGS OF THE NATIONAL MUSEUM 



BONELLIOPSIS, new genus 

Diagnosis. — Differing from Bonellia in having the nephrostome at 
the distal end of the nephridium instead of near the base, and in the 
possession of a much shorter presiphonal foregut; either left or right 
nephridium developed; females with two ventral setae; males without 
setae. 

Type, Bonelliopsis alaskana, new species. 

BONELUOPSIS ALASKANA, new species 

Figure 16; Plates 26, 27 

Description. — Body elongate, subcylindrical, blunt at both ends, 
20 to 65 mm. long, and commonly four or five times as long as thick; 
proboscis of usual Bonellia form seldom exceeding body length and 
usually considerably shorter. Contracted skin verrucose, the verru- 
cae squarish, not obviously larger in any particular region; when skin 
is stretched the verrucae flatten out into squarish glandular thicken- 
ings arranged in irregular longiseries. 




Figure 16. — Bonelliopsis alaskana, new species: Two males, the upper 1.27 mm., the lower 
1.9 mm. long, X SO. The spermatheca is most heavily shaded; the lightly stippled is 
parenchymatous and muscle tissue; sperm duct opens at anterior (right) end. 

Body wall thin, translucent. Inner, circular layer of muscles 
smooth, but in the region of foregut where the layer is thickest there 
is a division into slender fascicles. 

Setae 2, small, nearly straight, situated close together a short dis- 
tance behind mouth (4 mm. in specimen 44 mm. long). There is a 
short but broad interbasal muscle which usually presses upon the 
nerve cord and ventral blood vessel. 

Nepliridium 1; of six specimens dissected four had the left ne- 
phridium developed and two had the right. It is situated close to the 
nerve cord directly behind the setae. In some specimens the nephri- 
diopore is conspicuous externally. The nephrostome is conspicuous, 
terminal, with amply folded lips. When the nephridium is filled 



ECHIUROID WORMS OF NORTH PACIFIC — FISHER 253 

with eggs its tlistal oiul is usually iuvaginatt'd, coiicoaliiig the iicpliro- 
stomo (pi. 27, fig. 1). The mucosa is thrown into shallow longitudinal 
folds, most pronounced at proximal end. 

The two anal vesicles have the relative size and general form in- 
dicated in plate 26, figure 1, where only one is shown. Each opens 
by a small pore into the cloaca. The primary branches of the vesicle 
vary in number. In plate 2G, figure 2, is shown the tip of one of the 
major subdivisions including two of the smaller secondary branches, 
each of which bears several fumiels. 

Ali.ment.\ky c.\nal. As contrasted with Bonellia viridis the ali- 
mentary canal differs in having a much shorter forcgut, especially the 
portion between the gizzard and the intestine, corresponding in general 
to the "stomach" of Thalassema and allies. The abruptly enlarged 
intestine is produced forward into a coecum where the dorsal blood 
vessel envelops the intestinal wall. In the specimen of B. viridis that 
I dissected this was not differentiated (fig. 15). 

The pharynx is connected with the body wall by nmnerous radiating 
muscular frenula of which only a few are indicated in the figures. 
The mucosa is thick, verrucose, and not very different from that of 
the esophagus. The gizzard has stronger ring muscles than the 
esophagus (which shows a ringed structure), and the mucosa is 
thrown into consecutive ring folds. \Vhen spread out these ring 
folds subdivide into eight longitudinal divisions. Behind the gizzard 
the mucosa becomes al)ruptly tiiinncr, and between the gizzard and 
this elongated stomach there is a sort of pjdoric constriction, with a 
very narrow passage. The opening, on the stomach side, is surrounded 
by a fiange of tissue. There is no presiphonal ciliated groove, but a 
postsiphonal one runs along the intestine to the point where the 
ventral blood vessel and genital stolon join the hind-gut. The 
fecal pellets are slender blunt ellipsoids 2.5 mm. long. In one I found 
an ostracod and a small Balanus. 

Mesenteries. The mesenteries are continuous, very thin sheets, 
even in the postsiphonal region of the intestine. In plate 27, figure 2, 
an attempt has been made to show the principal mesenteries of the 
foregut, which are voluminous and folded when the animal is contracted. 
In this sernidiagrammatic drawing the left half of the animal only is 
slunvn. Wliat is probably the ventral mesentery (VAf) is attachcnl 
to body wall over a sinuous and not so smoothly regular course as 
shown in figure. It is fastened to the l(>ft sid(> of pharynx-esophagus, 
passing gradually to the side of the intestine oj)posite the siphon. 
The posterior part of dorsal blood vessel (/?') is attached to, or in- 
volved in, this mesentery. The dorsal mesentery (DAf) is attached 
to the body wall in a long spiral passing from right siile (dot and dash 
line) over to left side. Posteriorly it merges with the ventral mescn- 



254 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

tery on its left side, where the foregut enters the intestine. In the 
drawing the posterior face of dorsal mesentery is shown and only a 
small part of the anterior (or dorsal) side adjacent to pharynx. The 
attachment to the gut is along the lateral line, but behind gizzard the 
attachment moves over near to that of ventral mesentery. A special 
muscular mesentery indicated in plate 27, figure 1, but omitted from 
figure 2, joins the gizzard to the stomach. A special transverse dorsal 
muscular mesentery supports the dorsal blood vessel anteriorly. 
When the animal is extended the effect of these mesenteries is to form 
a somewhat spiral anterior cul-de-sac in which the nephridium lies 
and in which the eggs probably congregate. But when the neplnidium 
is full of eggs, as in plate 27, figure 1, there may still be hundreds of 
eggs, seemingly mature, in various parts of the coelom. 

Vascular system. The distribution of the principal trunks, 
shown in the figures, seems to be almost the same as in Bonellia 
viridis (fig. 15). The connection between the dorsal and ventral 
vessels is not direct, as in Echiurus, Ochetostoma, and Thalassema, 
but by means of lacunae in the intestinal wall, as is the case in 
Arhynchite. 

The gonad lying along the top of the ventral blood vessel, in the 
posterior half of body, seems to be identical with that of B. viridis. 

Male. The males are fomid in the foregut, from the pharynx to 
the gizzard, in the order of about a dozen to an individual. In one 
case I found two or three in the anterior part of the stomach. Dr. H. 
Heath, who collected the type series, examined several live specimens 
and found no males in the nephridium. 

The males vary from elongate-slender to the shortened state shown 
in figure 16. When fully extended they are more than twice as long 
as figure and only about half as thick. The sperm receptacle is 
situated posterior to middle of body and the duct opens at anterior 
end, or very close to it. There is very little free coelom, the body 
being filled with parenchymatous and muscle tissue, diagrammatically 
indicated by the more spaced dots in figure. No setae are present. 

Type.— U.S. ^M. No. 20603. 

Type locality. — Dutch Harbor, Unalaska, under rocks, at low 
tide; Harold Heath, August 1917, 15 specimens. 

Specimens examined. — In addition to the above, 25 specimens, 
without locality, U. S. National Museum. 

Remarks. — The type series was collected by Dr. Harold Heath, 
who found them in the intertidal zone under flat rocks. The worms 
were arranged around the periphery of the stone with the proboscis 
extended to the margin. In life the animals are light green, the 
color of "green prunes," which, in the contracted state, they some- 
what resemble. 



ECHIUKOID WORMS OF NORTH PACIFIC FISHER 255 

EUBONELLIA, new genus 

Diagnosis. — Bonclliidae with woll-dovolopcd l)ifurcate prol)oscis, 
110 setae, and a single (right) nephricUuni, the iHstal end of which is 
expanch^d mto the plicated rim of the large nephrostome; anal vesicles 
essentially as in Bonellia; male with sperm receptacle reaching nearly 
to posterior end of body; body wall thick. 

Type, Eubonellia valida, new species. 

Rrmarhs. — Tliis genus differs from Bonelliopsis, the only other 
known to have a strictly terminal nephrostome, in lackuig setae. 
Its foregut has an unusually extensive stomach or crop, between the 
gizzard and begnunng of siphon, this segment being short in Bonelli- 
opsis. The mesenteries of the mtestine of Eubonellia are m the form 
of strands, not a sheet. 

Parabonellia Onoda, '* based on Bonellia misakiensis Ikeda,'^ also 
lacks setae in both male and female. The nephridium has a small 
pedunculate funnel situated, not at the base as in Bonellia, but on the 
side of the vesicle near the terminal blind end. Ikcda did not show 
this in his figure of a "dissection" of the type, but it is figured by 
Onoda (fig. 1) and Sato '^ (fig. 2). None of these writers has figured or 
described significant details of the foregut, which are of importance in 
classification. 

The body wall of Parabonellia is thin whUe that of Eubonellia is 
very muscular, but this is a character varying with the contraction of 
the specimen and is scarcely of generic significance. 

In Eubonellia the male is about half the size of the male of Para- 
bonellia, which is described as nematodelike, 3.3 to 3.5 mm. long, and 
0.2 to 0.3 mm. in breadth. The sperm vesicle and canal occupy the 
anterior fifth to third of the body. In Eubonellia the vesicle extends 
nearly to the posterior end of the bodv, which is depressed, planarian- 
like. 

Profoboncllia Ikcda '* (type, P. mitsukurii, Sagami Bay, Japan, 300 
fathoms) has very strong setae with unusually well developed muscles. 
The proboscis is similar to that of ThahisKema, and the single (left) 
nephridium is similar to that of Bonellia. The nephrostome is a wide 
fimbriated funnel at the end of a slender tube opening into the nephrid- 
ium near the base of that organ. The anal vesicles aic similar to 
those of Bonellia rather than Ilamingia, which the proboscis might 
lead one to expect. The vascular system seems to possess a peculiarity 
in a rather direct connection between the dorsal vessel and the neuro- 
intestinal connective at a point immediately anterior (o the be- 
giiming of the siphon. Ikeda treats this very summarily in both 

"• On<xln 1«H, p. 418 (PtnidofiontlUa); 1035. p. HI (.Parabonfllia for PteudobontUia, prcocciiple<l). 
'• Ik<yU, 1W>I, p. 74, flKS. 24, 103-105. 
" ."^alo, 1935, p. 142, Ors. 1,2. 
'• Ikcda. 1808*. p. 259. figs. 1-4. 



256 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

description and figure, apparently not realizing that it is different 
from the relations of the two vessels in Bonellia. Neither his figure 
nor description gives any definite details concerning the foregut 
beyond the fact that it is very long. 

EUBONELLIA VALIDA. new species 

Plate 28 

Description. — The much-contracted type is oblong-cylindrical, 55 
mm. long, and about 20 mm. thick at middle. The conspicuous 
nephridiopore is 8 mm. behind mouth. Proboscis unusually broad 
and flat, terminally bifurcate, without a ventral sulcus. In its con- 
tracted state it is 30 mm. long and 7-9 mm. broad (pi. 28, fig. 3). 
The thick skin is thrown into annular welts, the furrows being fre- 
quently interrupted. These folds do not carry marked pustulate 
thickenings as in the case of Nellobia found at the same station. 
From the mouth a short narrow sulcus extends upon the constricted 
base of proboscis. 

Body wall tough and muscular, 1.5 to 2.5 mm. thick, the middle 
longitudinal layer being the thickest. The inner, circular layer is 
smooth, but somewhat fasciculated at anterior end of body. 

No setae or vestiges of seta sacs or muscles. 

The single large (right) nepln-idium has a terminal large nephrostome 
with voluminous lips. At the base a simple duct leads to the exterior 
approximately in the median line. 

The two anal vesicles are of the elongate dendritic type with a 
voluminous axial bladder having a few branches proximally. To the 
main stem and branches are attached singly or in clusters (pi. 28, 
fig. 2) the nephritic elements, which are characteristically very elongate, 
ending in pedunculate funnels. The vesicles are attached to each side 
of a very small cloacal bulb, the mucosa of which is thrown into longi- 
tudinal ridges. 

Alimentary canal. The anterior part of pharynx is attached to 
body wall by numerous frenula, but there is no peripharyngeal dia- 
phragm such as is characteristic of Nellobia. There is a rather delicate 
mesentery at the bend of the esophagus (not shown in figure). Food 
pellets form in the posterior part of esophagus (0^), as well as in the 
gizzard (60, and are collected in the elongate stomach. The wall of 
the stomach is almost transparent. Between it and the gizzard there 
is a powerful sphincter forming a sort of pylorus as in Bonelliopsis. 
There is a weaker sphincter between the esophagus and gizzard. 

There is no presiphonal ciliated groove. Much of the intestine is 
missing, but enough remains to show that there is an extensive por- 
tion, traversed by the siphon, which has thicker walls so that the 
pellets are not visible, whereas the postsiphonal intestine (with a 



ECHIIHOID WOKMS OF NORTH PACIFIC — FISHER 257 

ciliated grroove) has very thin walls and very compact pellets (j)!. 28, 
fi^. 4). The ciliated groove entls at entrance to small cloaca! bulb and 
a strand of tissue from the ventral blood vessel is attached at this 
point. 

The vascular system is of the usual bonelliid type and can be fol- 
lowed in the fi«rure. The ventral vessel divides into two at about the 
point where setae would be if present. 

The ^ronad, in the usual bonelliid position, is confined to the median 
third of the body and is inactive, there being: no sig:n of egg formation. 
In Xdlobia, dretlged at the same time, the gonads were active and the 
nephridium full of eggs. 

Male (pi. 28, fig. 5). One was found in the pharynx near mouth. 
It is possibly not fully matured; length 1.17 mm. The dark body is 
the spenn vesicle, the duct opening at the anterior end as in Bonellia 
nridis. The small spot back of the gonad is probably the excretory 
pore. Small indistinct masses of spermatozoa can be seen in the 
coelom, but have not been shown in the drawing. The gonad is situ- 
ated more posteriorly than in Bonellia viridis. 

Type—V.S.K.M. No. 20604. 

Type locality. — Albatross station 5021, Okhotsk Sea, off east coast 
of Sakhalin Island, lat. 48° 32' 30'' N., long. 145° 08' 45" E., 73 
fathoms, green mud, sand, pebbles, bottom temperature 30.9° F. 

NELLOBIA, '» new genus 

Diagnosis. — Bonelliidae without setae and possibly without probos- 
cis; one nephridium (.left), with a basal ncphrostome and swollen 
basal region, opening in the median line but without a genital groove; 
two compoand anal vesicles each consisting of numerous trees aris- 
ing from a sessile receptacle on either side of the very large muscular 
cloaca; terminal portion of hind-gut greatly enlarged; body wall very 
thick. 

Tvj)e, Nellobia eusoma, new species. 

Remarks. — This genus differs from Bonellia, Protobonellia, Para- 
bonellia, Eubonellia, and BoncUiopsis in the radically different struc- 
tun; of the aral vesicles and from all bonelliids in the extremely large 
muscular cloaca and enlarged terminal part of the hind-gut. The 
only group which hns fundamentally similar anal vcsi<'les is Acan- 
thohamln(}ia Ikeda ^ in which three trees arise indei)(!ndently on each 
side of the cloaca {A. ijimai) or more ninnerously as o cluster with 
probiibly some coiuiectif)n between the elements {A. shiplei). In 
Ilaminf/ia arctica Danielssen and Koren *' there is a very short (1 mm.) 



" Anneram of litmf.Uia. 

'^ Ikpiln, lOlO, p. l.Tfi, 1)1. 10; !kH> also lOOH, p. 01, pi. I. 

•1 Danlflsson an<l Korfn, IhHI, p. '.!n, [il 4 uml .'.. lU'". 1-1H. 



258 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

tube on each side of the cloaca to which very numerous small trees 
are attached at approximately the same point. 

The vascular system of Nellobia is similar to that of Bonellia and 
allies in the relation between the dorsal blood vessel and the neuro- 
intestinal trunk. No direct connection exists between the two by 
obvious anastomosis of terminal branches such as Danielssen and 
Koren figure (1881, pi. 5, fig. 14). 

NELLOBIA EUSOMA, new species 

Plates 29, 30 

Descrijption. — The single specimen is contracted to the maximum 
extent. The posterior end of the body is invaginated to form a cup- 
shaped depression . The intestine had been extruded, after the manner 
of holothurians, through a breach in the cloacal wall, and most of it 
is missing. 

Body of ^OTigZZm form without proboscis, 44 mm. long (allowing for 
posterior invagination), and 15-17 mm. thick at middle. Owing to 
contraction, the sldn is thrown into irregular transverse folds with 
frequent pustulate thickenings, less regular in the anterior ventral 
region (shown in pi. 29, fig. 1) than elsewhere. If the very short 
truncate snout is the remains of a longer proboscis, it is nevertheless 
covered with normal skin. The conspicuous opening of the nephrid- 
ium is close to the median line about 4 mm. behind mouth. 

The body wall is very muscular and in the contracted state about 
2 mm. thick. The middle longitudinal layer is the thickest, the 
inner circular layer the thinnest. The latter in the contracted state 
of the specimen shows definite fascicles of uneven width which would 
probably smooth out when the worm is expanded. 

The single neplmdium (pi. 30) is attached on the left of the nerve 
cord, but its duct passes under the cord to open in the median line. 
The nephrostome has very simple lips and passes into a bulbous and 
thick-walled proximal region. The distal compartment, filled with 
eggs, has the wall stretched to translucent thimiess but its proxunal 
constricted part has glandular walls furrowed longitudinally. There 
is a definite opening from the egg chamber into the proximal bulbous 
portion. 

The anal vesicles are peculiar. Instead of having a roughly treelike 
foi-m as in Bonellia and Bonelliopsis, the main vesicle is a sort of 
crescent-shaped pouch applied to each side of the large cloacal cavity 
and produced on the opposite or free border into numerous (a dozen 
or more) unequal dendritic subdivisions. The larger of these have a 
few main branches lilve a tree, which in turn are crowded with branch- 
lets (pi. 29, fig. 4) carrying many of the bulbous glandular elements 
ending each in a ciliated funnel. Around the base of these primary 



EClilUKOID WORMS OF NORTH PACIFIC — FISHER 259 

iirpliridia aro luiinorous siil)irlol>uIar umuiual yellow bodies arising 
from the l)ast' of the iiej)hritie eleiiieiits and from the wall of the collect- 
ing tube or branehlet. Xumerous much sni.'dicr brownish-yellow 
papillae occur on the walls of the main stems and branches. I 
could not find the opening into cloaca. 

Alimentary canal. The main features of the foregut are shown 
on plate 30. The pharyiLX is attached to body wall by very numerous 
crowded ratliating strands. This head cavity is separated from the 
rest of coelom by u translucent diaphragm (D) indicated in the drawing 
incompletely. Its central l)or(ler encircles the gut behind the pharynx, 
and what may be conventionally called the esophagus (extending to 
A' in drawing) has the muscular walls marked by prominent ring 
folds which cause the mucosa to be thrown into transverse welts. 
This muscle layer thins toward end of esophagus and the ring 
becomes narrower. The segment X-Y corresponds to the gizzard of 
BoncUiopsis. The annulation of muscle is closer. At Y the canal 
was broken, and it is possible that something was lost as the segment 
between Y and the beginning of siphon is very short. It corresponds 
to the so-called stomach of Bonelliopsv?. A tough mesentery unites 
loops of the esophagus, whereas in Bonelliopsis the thicker mesentery 
joins the gizzard to stomach. The esophagus has a continuous ventral 
mesentery attached ventrodextrally but the other mesenteric attach- 
ments to body wall are in strands or frenula. 

The cloaca is bulbous, with very numerous muscular strands uniting 
its rather muscular wall with l)od\' wall. The anterior of these strands 
pass between the branches of the anal vesicles. The very expanded 
hind-gut seems to be more than an accident of killing, as the condition 
of the mucosa indicates that th(> walls have not been unnaturally 
distended. The j)rominent ciliated groove continues from the narrow 
segment (all the rest of the intestine having been lost) to the beginning 
of the cloaea, where a strand from the ventral blood vessel ends. 
There is no intestinal coeeum at this point. 

The vascular system is of the Bonellia type. The neurointestinal 
connective (B^) spreads out fanwise where it joins the ventral vessel 
(B*) and its walls a[)[)ear to be glandular as if a i)art of the gonad 
complex. Actual ova are found iis far forward as the posterior border 
of this fan. 

The gonad is of the Bonellia type but extends uiuisually far forward. 

Male. Unknown. The foregut was (juite em[)ty, and no males 
were found in the nephridiopore. 

Typc.—\J.S.N.\[. No. 20005. 

Type locality. — Albatross station 5021, Okhotsk Seo, off cast coast 
of Sakhalin Island, lat. 48° 32' 30" N., long. 145° 08' 45" E., 73 
fathoms, green mud, sand, pebbles, bottom temperature 30.9° F. 



260 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 96 

Genus ACANTHOHAMINGIA Ikeda, emended 

Acanthohamingia Ikeda, 1910, p. 136. (Type, A. shiplei Ikeda.) 

Diagnosis. — Differing from Hamingia in having a well-marked 
genital slit extending forward toward mouth from the one or two 
nephridiopores, this containing, in two species, 8 or 10 very small 
setae imbedded in the skin; anal vesicles numerous, at least not in 
two symmetrical clumps which arise from a very short common duct; 
body wall thin; proboscis (when known) similar to that of Thalassema; 
males with or without ventral setae. 

Remarks. — The new species described below has necessitated an 
emendation of the original diagnosis since there are no minute setao 
in the genital groove. This groove, which extends forward from the 
nephridiopore, or pores, occurs in the three known species ^^ and is not 
present in any other genus. The general habit of the three species is 
much the same, as they are all of delicate build. The body wall is 
thin, translucent when expanded, and skin papillae are poorly devel- 
oped. The anal trees exhibit differences in the three species, being 
most alike in A. ijimai and A. paradola. These are numerous, inde- 
pendent, or semi-independent branched tubules, and differ from the 
condition in any other genus except Nellobia. But in Nellobia eusoma, 
which lacks any trace of a genital groove and is one of the most heavily 
built of all bonelliids, the anal trees spring from a bladderlike struc- 
ture applied to each side of the very muscular cloaca. The rudi- 
mentary bladder figured for A. paradola (left side) may well indicate 
the last trace of a similar structure. 

In A. shiplei and A. ijimai the male is long and slender and Iscks 
setae, whereas in A. paradola the male is lanceolate and planarian- 
like and is provided with two curved setae. 

ACANTHOHAMINGIA PARADOLA, new species 

Plates 31, 32 

Diagnosis. — Dijffering from A. shiplei and A. ijimai in the absence 
of minute setae from the genital groove of female and in the presence 
of a pair of ventral curved setae in the male; nephridia 2 instead of 1; 
anal trees numerous, slender, sparsely branched, arising for the most 
part independently from the very thin wall of the cloaca. Length of 
paratype 90 mm. (pi, 31, fig. 1). Color, pale flesh when seen on a 
white background. 

Description. — The general habit is much like that of A. ijimai but 
proboscis is lacking. The body wall is very thin and translucent, 
this thinness being accentuated by inflation. Along the midventral 
line the extremely slender nerve cord can be easily seen. The skin 



I A. shiplei Ikeda 1910, p. 136, pi. 10; Hamingia ijimai Ikeda, 1908b, p. 62, pi. 1. 



ECHIUROID WORMS OF NORTH PACIFIC — FISHER 261 

is marked by flat circular spaced spots slightly less translucent than 
the intervals. Back of the mouth is the characteristic genital groove, 
which normally is probably very narrow as in the type (pi. 31, fig. 3) 
but in the paratype (pi. 31, figs. 2, 2a) is spread apart owing to 
stretching of l)ody wall. In this groove are four males. In the type 
there is at least one. At the posterior end of groove are the openings 
of the nephridia: two in the type and another specimen (pi. 31, figs. 
4, 4(7); one in the paratype (pi. 31, figs. 1, 2). The hemispherical 
papilla sho\\Ti in these figures beside the nephridiopore is an artifact. 
Although there is but one opening there are two large iiepliridia 
containing eggs. 

As slated above, the body wall is very thin, on the order of 0.15 to 
0.2 mm. thick, and a single thickness is so transparent that printing 
can be easily read through it. The fibers of the longitudinal and 
circular muscles can be seen under magnification, but there are no 
bundles. 

All three specimens are in poor condition internally as the midgut 
and hind-gut are badly macerated, the contained pellets being adrift 
in the coelom. The more essential foregut can be made out with the 
associated blood vessels. 

The nephridia, in good condition, number two in all three speci- 
mens. They have exceedingly thin walls, and the small nephrostome 
and its short stalk are situated at the base, close to the swollen ducts 
leading to nepliridiopores. In the paratype the nephridia are about 
twice as large as those shown on plate 32. 

The significant features of the alimentary canal can be seen on 
plate 32. The pharynx is much inflated (but probably unnaturally) 
and has very thin walls. The esophagus can be traced to 0' where 
the fine longitudinal ridges of the mucosa change to equally small 
rings. Then follows an exceptionally long segment of the foregut 
in which I can find no marked division into gizzard and stomach (or 
"crop"). In this the contents are shaped into oblong pellets 1.5 to 
2 mm. long. Extremely slender and numerous frcnula connect the 
pliarynx to body wall. The continuous ventral mesentery of foregut 
is delicate and transparent. The position of the coils of foregut in 
figure has no significance as they had mostly broken moorings. All 
the foregut is thin-walled. 

In connection witli the vascular system the very considerable 
length of gut between attachment of dorsal vessel (/?') and ncuroin- 
testinal connective {IP) may be noted. The ventral expansion of 
IP is considerably farther back in another sj)ecimen. In keejMng with 
tiio rest of the animal the blood vessels arc delicate and the ventral 
trunk is very inconspicuous. There are numerous opaque noduk'S 
on the neurointestinal trunk. 



262 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

The nerve cord is the slenderest I have seen in a mature echiuroid, 
being only 0.135 mm. in diameter. 

The gonad, on the irregular margin of a mesentery dorsal to nerve 
cord, is invisible except under high magnification, as the cells are 
small and inactive. The whitish eggs in the nephridia are 0.5 to 
0.6 mm. in diameter. 

Anal vesicles are in the form of numerous slender tubes with spaced 
short lateral branches (pi. 31, fig. 6). These tubes are involved in 
and fastened to body wall by transparent but strong frenula. In the 
type nearly all the tubes of the left side spring from a small irregular 
common chamber closely appressed to the transparent wall of the 
cloaca, but oh the right side no such common chamber is present. 
Here the vesicular tubes arise independently from the cloacal wall. 
It is probable that the number of tubes increases with age. The 
ultimate subdivisions are not well enough preserved for exact delinea- 
tion but they resemble in general those of A. ijimai. Although the 
cloaca is not perfect in any of the specimens, that of the type shows a 
rather large subspherical chamber with transparent walls joined to the 
body wall by a multitude of frenula. 

Male. Males occur in the genital groove as shown in plate 31, 
figure 2. Probably the groove is normally as in the type, which has 
one or two males in it. They are depressed lanceolate in form, 1.2 
mm. long, and the tube of the sperm receptacle opens at the middle 
of the anterior end, the receptacle itself being about in the middle of 
body (pi. 31, fig. 5). 

TV^e.— U.S.N.M. No. 20601. 

Type locality. — Albatross station 4942, Kagoshima Gulf, Japan, 118 
fathoms, brown mud, black specks, bottom temperature 59.8° F., 
2 specimens. 

Specimens examined. — The above and 1 specimen (paratype) from 
station 4940, same locaHty, 115 fathoms (pi. 31, fig. 1). 

Xenopneusta, new order 

No blood-vascular system, the coelomic fluid being heavily charged 
with large blood corpuscles containing hemoglobin or hemoglobin 
and hematin; intestine with terminal portion in front of cloaca en- 
larged, thm-walled, functioning as an organ of respiration in connection 
with anus and cloaca. 

Family URECHIDAE Fisher and MacGinitie, 1928 

Diagnosis. — Differing from other Echiuroidea in the absence of a 
blood-vascular system, the corpuscles (red or brown in color from 
hemoglobin or hemoglobin plus hematin) free in the coelomic 
fluid; distal portion of midgut greatly enlarged and in connection 
with cloaca serving as a respiratory apparatus; foregut very long, 



ECHIUROID WORMS OF NORTH PACIFIC — FISHER 263 

includinE: a lonf;: frizzard botwoon an anterior lonp: crop and a posterior 
long stoniacli; proboscis reducetl to a scoop-sha})ed ii])per lip. 

Genus URECHIS Seitz 

Urechis Seitz, 1907, p. 352 (type, Echiiirus chilensis Max Miiller, 1852). — Fisher 

and MacGinitik, 1928a, p. 200. 
Spiroctetor Skorikov, 1909, p. 77 (typo, Echiurus unicincius von Drasche). 

Diagnosis. — Cylindrical or sausage-shaped echiuroids with characters 
of family. Body wall is very muscular, consisting of outer and inner 
circular layers and middle longitudinal layer, the latter the thickest; 
inner layer showing a fasciculate arrangement superficially. In the 
region of the posterior pair of nephridia is a zone of compound slime- 
net glands lodged in the verrucae of the skin. There are two or three 
pairs of nephridia, the basal nephrostome of which has long spirally 
coiled ciliated lips for collection of mature germ cells. The two an- 
terior setae have a strong interbasal muscle; one ring of curved anal 
setae interrupted ventrally. Traversing the coelomic cavity in front 
of the anterior setae are paired dorsoventral muscles (pi. 34, fig. 2, 
13). The alimentary canal has a definite pattern of attachment to 
body wall by muscular mesenteries, difTering in minor details in the 
three species (pi. 35, fig. 1). The slender foregut is very extensive, 
consisting of pharynx, esophagus, crop (subtended by a powerful 
muscular mesentery not attached to body), a long gizzard, and a 
stomach attached posteriorly by a strong mesentery. The greater 
part of the very long midgut is accompanied by the siphon, which 
starts close to distal end of stomach. The ciliated groove of midgut, 
which parallels the siphon, extends be3'ond it to the point where the 
gut is suddenl}' expanded mto the inflatable respiratory portion. The 
external ridge marking the ciliated groove continues along dorsal 
side of this expansion, passing distally to the right where it afi'ords 
attachment for mesenteries, but there is no groove inside corresj)ond- 
ing to it. This inflatable so-called "hind-gut" is equivalent to the 
terminal part of the midgut of Kchiurus (wliich is not enlarged). 
The only similar abrupt enlargement occurs in Nellobia eusoina. The 
true hind-gut or cloacal cavity is separated from the foregoing by a 
definite s{)liincter constriction and consists of a thin-walled ant(>rior 
liortion and a thicker-walled terminal section willi rugose mucosa. 
Verj' numerous frenula attach cloaca to body wall and account for 
tlie exj)ansion of cavity l)y which water is inhaled din-ing respiration. 
The anal vesicles are voluminous, slender sacs, always deflated, which 
open ventrally into terminal portion of cloaca. The glandular walls 
are externally rather cauliflowerlike and the entire inner surface is 
intricately plicated. The scattered ciliated funnels arc veiy tiny. 
The anus is eccentric to the circle of setae, being slightly nearer the 
ventral side fpl. 33, fig. 4). 

670329 — (6 4 



264 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 96 

Remarks. — Three well-known species have a very uniform outer 
facies. Although unicinctus of Japan has only two pairs of nephridia, 
it is indistinguishable by external features from small examples of 
caupo. The details of skin and setae are practically identical. The 
alimentary canal, except for minor details of mesenteries, is closely 
similar in the two species. Apparently umcindus does not reach the 
large size of caupo and chilensis, both of which have a third (posterior) 
pair of nephi'idia. The anterior setae of chilensis are blunt and 
scarcely tapered while in the other two species they are strongly tapered 
and sharp. In chilensis, as compared to caupo, dorsoventral muscle 13 
is weaker, the crop and gizzard are longer, and the interval between 
end of stomach and beginning of siphon is about three times greater. 
The attachment of the anterior end of the respiratory portion of 
midgut (pi. 36, figs. 1, 2) presents important differences. 

Urechis chilensis (Miiller), synonym U. farcimen (Baird), is found 
at Sandy Point, Strait of Magellan. My specimen, taken by the 
Hassler Expedition, was kindly donated by the Museum of Com- 
parative Zoology. 

A fourth species, Urechis novae-zelandiae (Dendy), awaits detailed 
investigation (Dendy, 1898; Poche, 1920). 

Dr. Carlos E. Porter, of Santiago, Chile, has called attention {in 
lift.) to the name Pinuca edulis Claudio Gay (1854, p. 475). Dr. W. 
L. Schmitt, to whom Dr. Porter sent the information, had photo- 
graphs made of the five pages covering the section on "Sipunculides" 
in the "Historia . . . de Chile." Four of these are pages 53-56 of volume 
3 published in 1849 and list Sipunculus lagena and S. cylindricus. 
Pinuca edulis is described in supplementary volume 8, published in 
1854, as follows: 

Afiade tomo III, pdg 56. Pinuca edulis. Por haber perdido los ejemplares 
que teniamos de este singular Sipunculiano, es precise d lo menos senalarlo & la 
atencion de los naturalistas y viajeros. Segun nuestro diario es de un bianco 
pardusco siicio y tiene de dos d. tres pulgadas de largo y como una de ancho. Su 
cuerpo es subcilfndrico, ligeramente hinchado en el medio y adelgazado en anabas 
puntas, siendo la anterior mucho mas obtusa que la posterior. El cuero es grueso, 
coridceo, un tanto arrugado en al traves, 16 que proviene de la reunion de una 
infinidad de pentitos mas 6 menos prominentes. La boca es pequena, arugada, 
rodeada, d poca distancia, de muy pequenos aguijones apenas visibles, subre- 
tractiles y dispuestos en circulo. El ano se halla d la otra extremidad y es 
bastante grande, liso, circular y un poco hendido. Un diseiio hecho en el lugar, 
senala hdcia el medio una reunion de pequenos cuerpos dispuestos en una banda 
circular de una linea poco mas 6 menos de ancho. 

Este animal que los habitantes comen cocido en la brasa despues de haberle 
quitado las dos extremidades, se halla en las arenas de la isla de Chiloe cerca de 
Castro, etc. En mi diario hallo notado que hace el pasaje de los Priapos d los 
Sipunculos. 

Without specimens from Chilo6 Island it is not possible to determine 
whether Pinuca edulis is the same animal as Urechis chilensis, which 
has not been reported so far north. If Pinuca is an Urechis it is 



FXIIIUHOID WORMS OF NORTH PACIFIC — FISHER 265 

obvious that Gay has the mouth and anus confused; and the h'ngth of 
2 or 3 inches is small for the average size. There are other discrep- 
ancies. In a Lrcchis 3 inches long, the anal setae are not "scarcely 
visible" but are conspicuous; no mention is made of the prominent 
anterior setae; no specimen of Urcchis displays "hdcia el medio inia 
reunion de pequefios cuerpos dispuestos en una banda circular de una 
linea poco 6 menos de ancho," whether "medio" refers to the body 
or to the anus (mouth), which just precedes this sentence in the 
description. 

The curious respiratory mechanism of Urechis, in connection with 
the loss of its blood vessels and the complexity of its blood physiology, 
relegates the genus to a very isolated position. Not less important is 
the behavior pattern whereby the requirements of respiration and 
food are beautifully met and coordinated. At every point specializa- 
tion of habit is matched by structural and physiological adjustment of 
the most delicate and efficient description. Along with this specializa- 
tion and complexity is an amazing viability best expressed by the 
term "tough." ^3 

All signs point to Urechis as being the last of a very ancient stock, 
one that maj' have flowered mto many species during Paleozoic times. 
It belongs to the honorable company of Lingula and those other 
aristocrats sometimes refeiTcd to as "living fossils." 

UnECHIS CAUPO Fteher and MacGlnitie 

F1GCHE8 17-19; Plates 33-35; Plate 36, Figures 2, 4; Plate 37 

Echiunia sp. Joh.nson and Snook, 1927, p. 178, fig. 153. 

Urechis caupo Fisher and MacGinitie, 1928a, p. 200, pi. 9, figs. 1-6; 1928b, 

p. 204, figs. 1-3, pi. 10. — Baumbercer and Michaelis, 1931, p. 417. — 

Reufield and Florkin, 1931, p. 185.— Hall, 1931, p. 400.— Sato, 1931, p. 

178.— Newby, 1932, p. 387; 1940; 1941, p. 303.— MacGinitie, 1935a, p. 341; 

1935b, p. 002; 1935c, p. 483; 1938, j). 208. 

Description. — The species reaches a large size. One specimen col- 
lected by G. E. MacGinitie at Humboldt Bay, Calif., measures 470 
nmi. long by 55 mm. thick. The largest specnnen from Elkhorn 
Slough, Monterey Bay, Calif., was 500 mm. long when fully relaxed 
in anesthesia, but after preservation it shrank to 375 mm. in length 
i>y 35 mm. in diameter. These were undoubtedly very old individuals. 
Average specimens are 150 to 180 mm. long. 

The surface of the body is traversed by fine irregular channels giv- 
ing a rugose api)earance, which is most pronounced in the head region 
anterior to the zone of slime glands. The latter, a sort of clitellum, 
is usually distinguishable by the circular trend of its fine furrows. 
Its anterior border coijicides roughly with the second j)air of nephiidia, 

n Sc« Rc'indfl and Florkin, 1031; nniimNrttcr nnd MIcharlLt, 1011; Hall, 1931; MncOlriitip, 103.15, 1935c; 
Newhy, IWO. IWl; Fisher and MncOlnltlo, 192«a. 



266 



PROCEEDINGS OF THE NATIONAL MUSEUM 



while the posterior border is spaced behind the third pair a distance 
about equal to interval between second and third pairs. This zone 
contains the slime-net glands. ''Three to 10 or even more glands 
are located on the outer surface and in the outer part of the sides of 
the papillae. These glands develop from the surface epithelium 
and are formed by an invagination of this layer of cells and the out- 
lying cuticle. The gland cells develop numerous terminal cones 
which penetrate into the cuticle of the duct (formed by invagination) . 
During the secretion of any one slime-net only about one-eighth of 
the gland cells are active. In these cells the terminal cones perforate 
the cuticle and open into the duct as minute tubules from which the 
fibrous secretion which forms the slime-net is discharged. The net 




Figure 17. — Urechis caupo Fisher and MacGintie: A vertical section of a mature slime- 
net gland taken while actively secreting; drawing by W. W. Newby (1941). 



ECHIUKOID WUKMS 01" XOKTH PACIFIC — FISHER 



267 



itself is i)robal)ly fibrous in iiaturo, althoutrli this lias not been demon- 
strated. (Figs. 17. IS.l 

"The body wall consists of the cutich*, the surface epitheliinn or 
epidermis, and the underlyin<; connective tissue called the cutis 
(Jameson, 1S99. p. 572) or corium (Seitz. 1907, p. 326). Tliese three 
layers constitute the skin. Beneath tlicni are the outer circular, 
the longitudinal, the inner circular muscle layers and the parietal 
peritoneum." -* 



inactive cell-/- 



active 




cell • ■ / 



Figure 18. — Urechis caupo Fisher and MacGinitic: A vertical section of the middle part 
of a slimc-nct gland taken while actively secreting; drawing by W. W. Ncwby (1V41). 

Anterior setae terminally tapered, sharp, curvetl, situated, in large 
specimens, 3 to 5 mm. !)ack of the groove leading to moutli and about 
the same distanec apart. They are metallic yellow, brownish at tip, 
8.5 to 10.5 mm. long. The flattened, curved exserted portion is more 
tapered and sharper than in chilcnsvi. A strong interbasal muscle is 

** Npwl.y, IMI, pp. 304. 3I.^; flRS. 1-10. Dr. Ncwby has subJprto<l the sllmo RlnnMs to a lliorouKh histo 
loKlcnl <itu'ly. He has kindly contributed tho orlRtnal drnwinKS of ORurct 17 and IN, which arc from the 
alKive paper. 



268 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

present, and numerous somewhat variable muscles radiate from the 
coelomic end. Inside the seta sac a short substitute seta is often 
present close beside the functional one. 

Anal setae curved terminally, sharp, 10 or 11 in number, and the 
dorsal are longer (8.5 mm.) than the ventral (7.3 mm.). The dorsal 
are the only ones used to any extent in cleaning out the burrow. 
When digging, the posterior end of the body is bent sharply forward, 
underneath, so that the ventral setae touch the ventral surface of the 
worm, while the strongly exserted dorsal bristles scrape the mud 
backward as the body is again straightened. This habit helps to 
explain the absence of a midventral seta, there being a broader gap 
at that point. The anus is eccentric to the circle of setae, being 
nearer to the ventral side (center of anus 6 or 6.5 mm. from dorsal 
setae and 4 or 4.5 mm. from the ventral). All bristles show conspic- 
uous cross-banding. 

There are three pairs of nephridia varying greatly in size according 
to degree of distension with eggs or sperm. In one specimen ex- 
amined the posterior tubes were 150 mm. long and 10 mm. in diam- 
eter, reaching two-thirds the total length of animal. The anterior 
pair is situated close to the setae. Rarely, one nephridium of this 
pair is missing. The nephrostome is on the anterior side at the base 
and the grooved ciliated lips are very long and spirally coiled. Mac- 
Ginitie (1935a) has shown that the superficial groove, V-shape in sec- 
tion in the male and more C-shape in the female, communicates by 
a slit along its bottom with what is virtually an almost closed duct 
or tube underneath. In both upper and deeper parts of groove the 
cilia beat toward the nephridium while on the outside of the lips, 
bordering the superficial groove, they carry materials in the opposite 
direction and incidentally help in the circulation of the coelomic 
fluid and contained blood cells. "As the eggs, blood cells, and other 
coelomic materials pass along the outside of the thread the eggs are 
caught in the external portion of the groove, are fed into the inner 
channel and then proceed to the opening leading into the storage 
reservoir [nephridium]. They are carried toward the reservoir at the 
approximate rate of 7 spirals per minute. Thus the eggs are sepa- 
rated from all other coelomic materials. As the eggs pass through 
the slit between the external portion of the groove to the inner chan- 
nel, they are under considerable pressure. They enter as a wedge, 
then become disc-shaped in the slit, and finally round out in the in- 
ner channel. Only mature eggs with indentations are collected in 
this way. As the eggs pass along the inner groove toward the storage 
organ they become oriented with the convex surface of one egg 
pushed into the indentation of the egg ahead, thus forming a com- 
pact chain. Blood cells (which range from 0.014 to 0.02 mm. in di- 
ameter) and immature egg cells pass along the collecting threads (i. e., 



ECHIUROID WORMS OF NORTH PACIFIC — FISHER 2G9 

tho spiral lips) without lodc:iii<]: in the extornal groove. Sperm is col- 
lected ill the outer groove of male collecting threads, fed into the 
iimer groove, and carried to the storage organ." " 

There is no permanent gonad. "Sex cells in all stages of develop- 
ment, from very immature ones to those wliich appear to be fully 
mature ore found in the coelomic fluid at all times of the year. In 
the case of the male, the apparently mature, free-floating sex cells are 
known to be functional." (MacGinitie, 1935c, p. 485.) "I have 
examined a male collected in winter as well as several specimens of 
both sexes collected in the summer and in these I could establish 
neither qualitative nor quantitative dilFerenccs in the sex cells at tho 
two seasons. Furthermore, in neither season did I find mitotic fig- 
ures in any area of the peritoneum nor could I establish any evidence 
of division by any of the cells which were free in the coelom. Thus 
there is no evidence in regard to the origin of the sex cells in Urechis." 
(Xewby, 1940, p. 7.) 

MacGinitie (1935a, p. 342) estimates that there are nearly 3 billion 
sperms present in the nephridia of an average-sized male and over 6 
million eggs in an equal-sized female. The eggs are 0.115 to 0.12 mm. 
in diameter according to MacGinitie (southern California specimens). 
C. V. Taylor measured 303 from the Monterey Bay region and found 
them to range between 0.123 and 0.144 mm. in diameter.^* The egg is 
ver}' clear, with a large nucleus, containing a nucleolus 0.012 to 0.016 
mm. in diameter. 

The anal vesicles, contracted, have a cauliflower surface beset with 
minute ciliated funnels. They empty, vcntrally, into the posterior 
part of the cloaca. 

In addition to abundant sex cells, the coelomic fluid is filled with 
nucleated red or brown blood corpuscles subcircular in shape and up- 
ward of 0.035 mm. in diameter, together with very numerous amoe- 
i>oid cells, yellowi.sh when aggregated. "The color of the blood 
varies from the i)urest oxyhemoglobin red to the darkest brown-black 
or a blacidike Chinese ink, even after complete saturation with o.xygen. 
The red color is due to liemoglobin homogeneously distributed within 
tho blood cells. Wiieiiever the color is brown, besides this hemoglol)in 
there is another granular pigment of brown color within the cells 
wliich will be proved to be hematin. Red blood was encountered in 
some few of the smallest indiviihials and in some of the very largest 
sex-mature females. The majority of the individuals, of medium size, 
contained brown or brown-bhick blood." (Baumberger and 
Miehaelis, 1931, p. 417.) 



»» Mnrfilnltlp, llO.'*, p. 340. A carrful pnpcr bawd upon observation of living material and tho only one 
dcsrrlMnit tho l)»>hii%ior of the nophrldlnl nppcndoKos. 
» Pbyslologlcal Zoology, vol. 4, p. 430, 1031. 



270 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

Alimentary canal.^^ — As indicated in the family and generic 
diagnoses, the most characteristic f eatm-es of the aUmentary tract are 
the extremely long presiphonal foregut, with its very extended gizzard 
segment, and the ''hind-gut" modified to act as a respiratory organ. 
These features are closely similar in the three species, all of which I 
have dissected. The apparent differences in the published figures 
are due to limitations of material or faulty observation. In plate 34, 
figure 1, the greater part of the siphonal and part of the postsiphonal 
gut have been removed, A very contracted specimen was used owing 
.to limitations of plate. Here it is obvious that the foregut is con- 
siderably longer than the body. In a well-expanded preserved speci- 
men, 300 mm. long, the pharynx is 30 mm. long; esophagus 40 mm.; 
crop 85 mm. ; gizzard 85 mm. ; stomach 50 mm. ; total 310 mm. In an 
expanded specimen the distance between the position of the stomach 
and anus may equal nearly one-third body length and the muscular 
mesentery may be, as in the specimen under discussion, 35 mm. long. 
The posterior attachment of this mesentery is indicated also in plate 
36, figure 4, M\ 

The pharyngeal lining is thrown into very coarse longitudinal 
folds, which really begin on the ventral side of proboscis and run 
directly to the esophagus, diminishing in size. In the esophagus 
the much smaller ridges are cross cut by deep narrow channels, which 
divide them into rings of oblong verrucae, giving the exterior a ringed 
appearance. The esophagus is definitely begun at the last of the 
dorsolateral mesenteries of pharynx (pi. 34, fig. 2, 15). In this 
figure an attempt is made to show the muscular mesenteries of the 
pharynx by a view looking forward into the head region from just 
back of the first pair of neplnidia. The pharynx has been pulled 
upward by the contraction of these dorsal and dorsolateral mesenteries 
1-7 (left side). The ventral mesenteries {IJj) are the least variable. 
Muscle 13 (paired) is not connected with the alimentary canal but is 
attached below to body wall in front of the seta, and above, dorso- 
laterally. All these muscular mesenteries are characteristic of the 
genus and on direct comparison appear to be more robust in chilensis 
except 13, which is better developed in caupo. 

The crop (crop 1 of Seitz) is subtended by a strongly muscular 
mesentery not attached to body wall. The posterior part of esoph- 
agus (pi. 34, fig. 1, X) loses its ringed appearance, the mucosa hav- 
ing deep, fine, longitudinal folds. In the crop the mucosa is again 
regularly verrucose, but of finer texture than in esophagus. In the 
gizzard the mucosa is thrown into strong ring folds, formed by the 
annulate muscles of gizzard wall, conspicuous superficially. The 



" Embleton, 1900, pi. 8, gives some figures of the histology of alimentary canal of U. unicinctus; Seitz, 
1907, pi. 31, for U. chilensis. 



ECHIUROID WORMS OF NOKTH PACIFIC — FISHER 271 

crop and gizzard of chilcnsis by direct comparison are definilcly 
longer than in caupo by about 25 percent. 

The stomach (crop 2 of Scitz) corresponds to that of Thalasscma, 
and the mucosa lias about 12 distinct longiseries of compressed ver- 
rucac. Externally the stomach has the longitudinal zonation charac- 
teristic of Thalassema and allies. It ends abruptly at the beginning 
of the much wider midgut with its ciliated groove. About 3 mm. from 
end of stomach the siphon begins, while the ciliated groove continues 
adjacent to it, along the inside of intestine (pi. 35, fig. 5). A strong 
muscular mesentery subtends the stomach and is attached posteriorly 
a little to left of nerve cord {^P). In chilcnsis the siphon begins 9 mm. 
from end of stomach as compared to 2.5-3 mm. in caupo. 

The course of the intestine in a fully expanded specimen is shown 
in plate 35, figure 1. The anterior and posterior portions of the body 
are omitted. Plate 36, figure 4, shows the cloacal region of the same 
specimen. It will be seen that the siphonal part of the intestine is 
very long and includes two anterior and two posterior bends. Three 
segments of the gut are attached by muscular mesenteries on the right 
side of body and three (including the big "hind-gut") are attached 
on the left side. The foregut is not attached to body wall except by 
the strong phamygeal mesenteries and the mesentery of the stomach 
(-IP). In this figure the breadth is accentuated because the body 
wall is pinned out flat. The mesenteries of siphonal gut farthest to 
right are attached about halfway between midventral and middorsal 
lines. 

The postsiphonal "small intestine" is rather short and is anchored 
by heavier mesenteries than are found on the siphonal portion anterior 
to the last loop. The "hind-gut," used as a respiratory organ, varies 
in dilation and consequent thickness of wall in difrerent specimens. 
The wall is usually thin and translucent. It is firmly anchored along 
its entire length on the left side of the nerve cord. On plate 36, 
figures 1-3, I have shown the attachment of the anterior end of the 
"hind-gut" in the three species. Unless the single specimen of 
cliilensis available for dissection is abnormal, there is considerable 
difference between it and caupo. 

The cloaca is probably normally elongate as shown in plate 36, 
figure 4. It is here shown opened for the entire lengtli. 'J'he mucosa 
of the posterior third is deeply furrowed longitudinMlly, and in this 
part, on the ventral side, are the openings of the two anal vesicles. 
The fecal pellets which sometimes crowd the i)ortion of small intestine 
shown in plate 34, figure 1, are cylindrical with rounded ends (pi. 35, 
fig. (V). 

T\\v fresh colors of the viscera in an anesthcti/cd specimen are: 
Foregut, pale flesh or skin color; anterior third of midgut i)ale gray- 
green mottled Nsith brown; middle third, mottled yellow antl dull 



272 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

gray-green; posterior third pale gray-green; siphon, pale jade green; 
respiratory gut, translucent raw sienna. 

T?/^g.— U.S.N.M. No. 19616. 

Type locality. — Elkhorn Slough, an estuary of Monterey Bay, 
Calif. ; shallow water, muddy sand. 

Distribution. — California: Humboldt Bay, Tomales Bay, Mon- 
terey Bay (see note below), Morro Bay, Newport Bay. With one 
exception all specimens have been found living under essentially 
estuarial conditions; that is, in quiet bays or sloughs in sandy mud. 
As a rule the openings of the burrows are under water at low tide, 
but are sometimes uncovered. However, in June 1923 I found one 
good-sized specimen in a bucket holding flounders and other fish 
caught in essentially open sea conditions near the Hopkins Marine 
Station, Pacific Grove, Cahf. This record points to the probable 
occurrence of the species at moderate depths almost anywhere off the 
coast of Cahfornia where mud of the proper consistency for permanent 
tunnels is present.^* 

History. — The first specimens of which I have any knowledge were 
collected in 1903 by C. S. Thompson, at Morro Bay, Calif., and 
brought to Stanford University. Some of these, in a good state of 
preservation, are still in the museum there. In 1920 I found one 
specimen in Elkhorn Slough, Monterey Bay, where a few years later 
Dr. Myrtle Johnson collected the examples from wliich the figures in 
"Seashore Animals of the Pacific Coast" were drawn. In 1923 a 
specimen was brought in by flounder fishermen from the sea bottom 
off the Hopkins Marine Station. It was not until 1926 and 1927, 
however, that the animal was studied. In connection with an 
ecological exploration of Elkhorn Slough, Prof. G. E. MacGinitie, 
then a graduate student working at the Hopldns Marine Station, 
found them in quantity. By means of narrow aquaria filled with mud 
("limoria") and glass-tube facsimiles of the actual burrows, he was 
able to observe living animals under essentially normal conditions, 
for Urechis seems to be insensible to light. Every important fact 
in the ecology of Urechis has been discovered by Professor Mac- 
Ginitie.^^ 

Habitat. — The first field studies were made at Elkhorn Slough, a 
shallow estuary, tributary to Monterey Bay, where the water, al- 
though slightly warmer than that of the ocean (which here varies 
from 49° to 57° F.), has practically the same salinity, there being 
usually a free interchange with each tide. In this inlet dwell a con- 
siderable variety of bivalves, some of which are much sought for food. 
There are two very interesting decapods, Callianassa californiensis 
Dana and Upogebia pugettensis (Dana) , which, like Urechis, construct 

" Dr. Earle H. Myers tells me he has found Urechis in the stomach of dogfish caught northwest of San 
Francisco Bay entrance (Golden Gate). 
» Fisher and MacGinitie, 1928b; MacGinitie, 1935b, pp. 682-686, 688, 715, 717; 1938, p. 208. 



ECHIUROID WORMS OF NORTH PACIFIC — FISHER 



273 



tumiels ill the mud tiiul conduct a more or less permanent manage. 
The mud teems with amiehds such as Lumbrinireis, and there are 
hterally acres that have a greenish tinge from the tentacles of Phoro- 
nopsi^ rirulis Hilton. Zostera grows in jiermanent patches and sup- 
ports a characteristic association of animals. At favorable times wide 
expanses support a growth of green Enteromorpha which, either fresh 
or decayed, is an important food element, shice the bulk of animal 
life consists of detritus feeders. 

At low water broad areas are left bare, but Urechis usually excavates 
its home where the entrances are not exposed at lowest tide. A few 
places were found where they are exposed at lowest tide. 




Figure 19. — A, Urechis caupo Fisher and MacGinitie, in resting posture. B, Position 
assumed while dig^'ing with anal setae. C, Plan of Urechis tunnel, the worm in situ 
pumping water through the slime-tube. Water enters at C, where there are two Cleve- 
landia ios (one outside); at a, Hespernoe lies in wait to feed on tube when it shall be swal- 
lowed; b, the tiny clam Cryptomya californica; c, Scleroplax; d, Clevelandia creating 
disturbance; e, eruption of mud cloud on ventilating current; castings around exit. 

The tunnel (fig. 19, C), never carried very deep, has two entrances 
and is in the form of a widely expanded U, of which the uprights are 
nearly perpendicular and the bottom horizontal. The apertures are 
small, being about one-third the diameter of the tunnel itself. Around 
one of the openijigs there is a considerable cjuantity of castings. The 
greatest distance between entrances measured 38 inches, the shortest 
16 ii\ches. Twenty-seven was the average for many measurements. 
The distance apart of the two entrances depends upon the size of the 
animal but not proportionately so, for small specimens have more 
extensive burrows for their size than larger ones. The largest speci- 
men obtained was 19.5 ijiches long when relaxed in anesthesia, and 
the smallest was 1 inch. 

The burrows have a permanent aspect and none of those contijiutiily 
observed was found changed except that occasionally one had a Jiew 
entrance. The animals grow very slowly, and so the enlargements need 



274 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

be made only at infrequent intervals and then only by widening the 
U and extending one of the entrances. 

Locomotion. — A Urechis can move along a smooth surface in much 
the same manner as an earthworm. It elongates the anterior part 
of the body and then forces forward the viscera and the water con- 
tained in the respiratory gut by contracting the posterior region and 
relaxing the anterior. When most of the body weight is in the anterior 
end the posterior portion is draAvn up. These movements are re- 
peated as the animal proceeds. Its method of locomotion within the 
burrow is quite similar except that the animal has the added advan- 
tage of being able to wedge the anterior portion against the sides of 
the burrow. Its rapidity of locomotion approximates that of an earth- 
worm. It can move considerably faster when in the burrow than 
when out of it, and it can move backward nearly as fast as forward. 

Digging. — When digging a tunnel Urechis forces its proboscis into 
the mud and works out a hole until the body can be drawn into it. 
This process is continued until the worm completes a U-shaped 
tunnel open to the surface at both ends, so that a supply of fresh 
water may be pumped through the tube by peristaltic movements of 
the body. Then the bore of the tunnel is enlarged by scraping ma- 
terial from the sides by means of the anterior setae, working it back- 
ward with the anal setae, and finally blowing it out the "back door" 
by a blast of accumulated respiration water from the hind-gut. To 
loosen sand from the sides of the burrow, the oral setae are protruded, 
then drawn backward through the sandy mud. This digging is done 
on all sides of the burrow as the animal can rotate its body at will. 
The setae are shed occasionally and renewed. 

The use of the anal setae, which form a ring of 10 or 11 a short 
distance from the anus, is higlily characteristic and was carefully 
observed. The sharp retractile bristles curve forward. The mid- 
ventral seta is lacking, ard the pair on each side are distinctly shorter 
than the four or five dorsal setae. Wlien a certain amount of loosened 
debris accumulates from the activities of the anterior bristles, Urechis 
crawls over it and forces it backward, in one of two ways: either by 
blowing the sand along with anal-water jets, augmented by the vigorous 
ventilating stream of the tube, or by turning under the posterior end 
of the body and then vigorously straightening it (fig. 19, B). The 
loosened material is shoved along the tunnel, whereupon the anal jet 
and ventilation current propel the finer detritus still farther. The 
animal backs up and repeats the process. When the posterior end is 
folded under, the dorsal setae are strongly everted and their forward 
curvature favors efficient scraping. The ventral setae (now dorsal 
in position) are against the vential body wall and do not function. 
A reason for the smaller ventral setae (and the absence of the mid- 



ECmUHOID WOHMS OF XOHTH PACIFIC — FISHER 275 

ventral sctji) is now apparent, if wo linve faith in the cfTicacy of use 
and nonuse in (Joterniining the rehitive size of siniihir orpins. 

Castings are sometimes ejected fioni the burrow by tliis flipping of 
the posterior end of the body (wliieh ean be aihnirably imitated with 
the forefinger), but usually only by water currents. Castings are 
allowed to accumulate and then are ejected in (juantity from one 
entrance. When digging downhill the animal shoves the soil along 
the body and then out by backing up the burrow, forcing the sand out 
the last inch or two by water currents. The opening then resembles 
a miniature vok-ano with line dark sand sj)outing out and the roily 
water trailing oil' from the crater like smoke. A major convulsion 
will carry out fragments of shells 2 or 3 mm. in section. Larger 
objects are avoided or allowed to fall toward the lower part of the 
burrow where the}' are buried. Doubling the velocity of water in- 
creases its carrying efficiency directly as the sixth power. 'J'hc nar- 
rowed mouth of the tunnel undoubtedly aids in increasing the force 
of these "volcanic" manifestations and hence their efficiency in re- 
moving sizable debris. 

Once Urechis is settled in a permanent home its daily activities 
consist of respiratory movements, obtaining food, cleaning the bur- 
row, and resting. 

Hespiratory 7noreme7tts. — There are two separate movements con- 
cerned with the respiration of Urechis: (1) The peristaltic movements 
along the bod\' which pump fresh water into the tunnel and move 
that within respirator}' chamber of the intestine; (2) the inhalations 
and exhalations, through the anus, for which the muscular cloacal 
chamber, resembling that of a holothurian, su|)i)lies the ciiief motive, 
power. 

The inhalations are from 1 to upward of 30 iji succession (witliout 
an exhalation). Exhalation is usually a single discharge although 
infrefjuently a rest may occur duiing a period of exhalation. The 
rote of breathing is not uniform. For instaiu'c, 2 inspirations cover- 
ing 25 seconds were followed by an expiration period of 10 seconds, 
whil(! in another instance 7 inspiiations occupied 2") seconds, the ex- 
piration 10; 24 hisj)irations occuj)ied 70 secojuls, the single expiration 
50; 30 inspirations occupied 90 seconds, the expirations only 25. 
Inspirations fewer than 12 predominate in a total of 11 cycles timed. 
These tinu's were tak(;n on a specimen lying in a pan of water. In 
its natural environnient Urechis bi-eath(!S more .'^lowly, but with the 
same irregidarily. 

The peristaltic movements of the body which serve to propel water 
thr(tugh the tul»(' are oven more erratic. The wa\'e, which expands 
the body to lill the burrow, begins at the base of the j)roboscis and 
passes along the body at varying rates for dilferent waves or even the 
same wave in diirereul parts of the body. As one wave arrives at the 



276 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

posterior portion of the body a new one begins at the anterior end, 
Urechis is normally always in control of the water and senses any- 
thing which may attempt to pass through the burrow. 

Feeding. — The unique method employed by Urechis to gather 
nourishment is generic and furnishes a striking example of the coordi- 
nation of adaptive structure and behavior. 

A short distance back of the oral setae is a zone of compound 
mucous glands, which form a sort of clitellum very faintly differen- 
tiated externally by the ringlike arrangement of the low rugosities of 
the skin. These glands are specializations of the simpler and more 
numerous multicellular flask-form mucous glands of the integument. 
The slime-net or girdle glands, as they have been called, secrete a 
fairly long transparent mucous tube, or funnel, whose upper, open 
end is fastened near the mouth of the tunnel while the lower remains 
attached to the clitellum. This strains from the ventilating, or respir- 
atory currents, all minute particles as the water flows through and 
when sufficiently loaded the tube is swaUowed. The process is 
repeated as long as the animal feeds. Urechis readily adopts a glass 
substitute for its normal burrow so that feeding reactions can be closely 
followed (pi. 37, flg. 1). 

Just before starting a tube, the body is constricted at the clitellum. 
This region is then expanded until it presses firmly against the sides 
of the burrow (usually near the mouth, but sometimes in the hori- 
zontal portion) with which it remains in contact for about 2 seconds. 
During the spinning process, which occupies only a few minutes, the 
constricted portion of the body anterior to the clitellum (whence the 
slime is issuing) undergoes a curious spiral peristalsis (pi. 37, fig. 5) 
easily detected by watching the nerve cord, which shows through the 
pink body wall, while back of the clitellum the normal respiratory, or 
pumping, peristalsis is taking place. 

The tubes vary in length from about 2 to 8 inches without apparent 
reason. As the tube lengthens Urechis backs down the tunnel, and on 
completion the spiral peristalsis anterior to the clitellum ceases, being 
replaced by a faint normal peristalsis, the main wave starting just back 
of the attachment of slime tube to the body. These normal ventilating 
reactions are kept up until the animal, apparently sensing the blocking 
of the water current by the clogging of the mucus with detritus, slips 
the tube forward "over its head." In doing this it deftly catches the 
hind edge of the tube by expanding the proboscis and bending it back- 
ward, collarwise, against the inflated nuchal region, until the muscular 
pharynx is able to pick up and suck in a portion of the margin (pi. 37, 
fig. 7). When diatom culture or detritus is introduced with a pipette 
the slime tube is soon swallowed; but if unmolested, Urechis may 
continue pumping for an hour before the tube is clogged. 



ECHIUROID WORMS OF NORTH PACIFIC — FISHER 277 

Usually only a few minutes are required for swallowing the tube, 
but the time depends upon length of tube and the amount of detritus 
intercepted. ^Yhen the tube has been swallowed up to the point of 
attachment the animal makes a movement to release it from the 
sides of the burrow similar to the reaction while digging with oral setae. 

The food funnel is porous to liquid but will intercept the smallest 
particles. Phenol red passes through everywhere, but no carmine 
particles ever do. Under the microscope no openings can be detected, 
but particles approaching a micron in diameter are lodged in the 
nmcus. 

When first secreted the tube is perfectly transparent, but as it 
collects detritus it becomes gray and its outlmes are easily seen. 
Peristalsis becomes more energetic as the tube-wall fills. 

When spuming the tube or lying at its lower end pumping water 
through it, Urechis is very sensitive to disturbances. If water is in- 
jected into the mouth of the tunnel, the animal immediately ceases 
movement and remains perfectly still for a minute, then slowly 
resumes peristalsis. If the disturbance is too great, it will drop out of 
the tube and retreat toward the center of the burrow, returning later 
to eat the slime tube. While it is lying at the end of a completed slime 
tube any sUght disturbance such as the introduction of a little mud or 
fresh clean meat will cause Urechis at once to pass the tube forward and 
begin swallowing. No large particles are ingested. They are rejected 
as the tube is being swallowed. 

Urechis feeds to some extent, although not very efficiently, when 
lying without its burrow in an aquarium. In such a position it will 
swallow sediment from the bottom of the aquarium gathering it with 
the proboscis. 

Resting. — After a period of feeding Urechis goes to the horizontal 
portion of the burrow, contracts its body so that it fits the tunnel 
snugly, and lies in a state of suspended activity during which even 
respiration ceases. These rests may last for an hour or more, but the 
long rests are always preceded by one or more short rests, which last 4 
to 8 minutes, and between which respiratoiy water is expclh'd and 
more taken in (fig. 19, A). 

Commensals (pi. 37, figs. 1-3). — Urechis has three pcrmaiK'nt 
commensals: A polynoid annelid, IIes])er()ii<>c adirnior (Skogsberg), 
and 2 pinnotherid crabs, ScleropUiT. (franv/uta Katlibun and Piniiixa 
Jranciscana Rathbun. Sometimes all three are found in (he same 
burrow, but usually only a Ilesperotioe and either a ScleropUix or a 
Pinnixa. In addition, the litth; Cryptomya californica (Conrad) 
projects its siphons into the burrow to make use of the water in the 
burrow for its source of food and oxygen. The goby Clevelnmlia ios 
(Jordan and Clilbcrt) uses the burrow as a retreat rather than a resi- 
dence, as the little fish freely forages outside, returning when alarmed 



278 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 9« 

or when the entrance is left exposed by low tide. On such occasions 
one to five gobies may be taken from the upper part of the tunnel. 
A goby left at the laboratory for several weeks in a glass burrow ap- 
peared contented. It would pass from one end to the other, wriggling 
past the Lrechis as if accustomed to doing so. At Newport Bay a 
pair of either Betaeus longidactylus Lockmgton or Crangon [Alphaeus] 
californiensis (Holmes) have been found permanently established in 
the burrows of U?^echis. The former is also recorded from Elkhorn 
Slough from Urechis burrows (MacGinitie, 1935b, p. 706). 

Hesperonoe adventor, which ranges in length when alive from 15 to 
50 mm,, is commensal with Lrechis throughout its range from Hum- 
boldt Bay to Newport Bay and normally is not found outside the 
burrows. The food of Hesperonoe consists of particles rejected by 
Lrechis when swallowing its slime tube. These particles consist of 
either living or dead animals which wash down the burrow with the 
current and become entrapped in the slime-net. Sometmies when 
Lrechis is swallowing its slime tube the polynoid will crawl forward and 
eat part of the tube and contents. It is very aggressive toward in- 
truders within the burrow other than the commensal crabs. Only one 
Hesperonoe occurs within each burrow, and if another enters the two 
will fight until one is killed or driven from the burrow. Other annelid 
worms wliich may find their way into the burrow are speedily dis- 
patched (by means of the short eversible toothed proboscis) and 
devoured. 

Hesperonoe rests with its dorsal surface in contact with the body of 
Lrechis, moving along the bm-row with the latter by making little 
short runs as the peristaltic movement of the body of Lrechis passes 
by. It always faces in the same du-ection as Lrechis, and when the 
latter turns in its burrow the annelid quickly does likewise. Hespero- 
noe is also commensal in Echiurus tunnels. 

Scleroplax ranges from 3 to 13 mm. across the carapace and is com- 
mensal also in the tubes of Callianassa calijorniensis and Upogebia 
pugettensis. It rests facing the side of the burrow, the chelipeds 
turned up in front and the last pair of legs raised behind. In this 
posture it can travel sidewise along the tube much faster than its host. 
Its food consists of particles which wash into the burrows or are un- 
covered by the hosts. Pinnixa franciscana screens detritus by 
means of its second maxillipeds, and it will also feed on particles of 
worms, clams, etc. Scleroplax has never been observed screening 
plankton. As many as six Scleroplax have been taken from one 
Lrechis burrow. A male and a female are often found together or two 
females. In one instance an ovigerous female was found with an 
ovigerous Pinnixa, and in another burrow a male Scleroplax and a 
male Pinnixa. 



ECHIUROID WORMS OF NORTH PACIFIC — FISHER 279 

Enemies. — Urechis piohahly attains a ripe old aj^o. Five spociinons 
of (lifToront sizo, kept in mud in the laboratory for over a year, appeat 
not to have p-own. However, as their food is prineipally detritus, and 
as natural conditions tire necessary to keep this stirred up in order that 
any quantity may be drawn into the slime net, laboratory <;rowtli tests 
are not convincini:. Yet what mitrlit be temied the settled habits of 
the creature ami the scarcity of very small specimens point stronjj:ly 
toward lonjicvity. The only animal known to prey upon them is the 
stinj: ray (Myliohattis califorjiicus Gill), which can dijj: out an occa- 
sional Urecfii.'^. In tiie ocean, however, small worms are possibly eaten 
by flatfishes, which repdarly feed upon Listriolohus pelodes. As 
already noteil, Dr. Earle H. Myers found Urechis in the stomach of 
small sharks. 

The period of mortality probably comes during the larval stajre. 
The small goby {Clevelandia ios) is extremelj' numerous, darting here 
and there, for any moving particles. These fish range from half an 
inch to IK inches in length and often devour objects so small as to be 
invisible to the observer. On one occasion 400 of these little gobies 
were netted from a hole, 3 by 6 feet, left by clam diggers. In addition 
the tiny Urechis must run the gauntlet of a host of small prcdaceous 
crustaceaiLs, annelids, nemerteans, and mollusks which forage on the 
surface and in the upper layers of mud. Once established in a biuTow 
Urechis is relativel}' safe. 

Parasite. — I have found rather numerous cestode larvae 0.25-0.32 
mm. long in the proximal end of the siphon where they perhaps cause 
the hernialike swellings of the siphon wall (pi. 35, figs. 4, 4a, 5). 
Probably the adult is to be found in the sting ray. 

Spawning. — Stored sex products are found in the ncphridia through- 
out the 3'ear. MacGinitie (1938, p. 208) states that normal spawning 
takes place during a short season, usually in spring or at the begimiing 
of summer as the temperature of the water rises. One male which he 
kept in the laboratory for two or three years spawned on May 24 and 
25. Just prior to spawning the worm came nearly to the opening of the 
glass tube which served as a habitation. Three welts were thrown around 
the body so that the circular creases were just anterior to each of the 
three pairs of gonopores, and the gonopores themselves were somewhat 
protruded and turned toward the anterior end of the body, and, 
therefore, t^)war<l the opening of the tube. The gonopores became 
quite consj)icuoijs; this was folh)wed by several retching movem(>nts, 
as if the aninud were attempting to n'giirgitate, and then sperm issued 
in a stream from each gonopore, VVIu-n the sperm ceased to bo 
expelled, the animal underwent violent jieristalsis, the waves running 
from tin- posterior to the anterior end, causing the sperm to i)our out 
of the glass tube. The retching, followed by the violent antiperistalsis, 
was performed three distinct times. On both (hiys after s[)awning the 

B70329 — 40 5 



280 PROCEEDINGS OF THE NATIONAL MUSEUM vol. se 

worm went back to the bottom of the tube, pmnped vigorously for 
§ome time, and then resumed feeding. During spawning the body of 
the worm was much more elongated than normally. The spawning 
on May 24 occurred at 4:30 p. m., that on the following days at 9:20 
a. m. Although tae nephridia were emptied the first day of spawning, 
the movements and procedure on the second day were the same as for 
the first spawning, but very little sperm was discharged. 

The embryology of Urechis caupo, outside the scope of this paper, 
has been thoroughly described and figured by Dr. W. W. Newby 
(1940). In this paper, which merits the highest praise, the relation 
of the Echiuroidea to other phyla is fully discussed. 



ADDENDUM 



In 1942 Dr. Sixten Bock published an important memoir "On 
the Structure and Affinities of * Thalassema' lankesteri Herdman and 
the Classification of the Group Echiuroidea." Owing to delays 
occasioned by the war, it has been possible to incorporate only the 
most important systematic data in the foregoing report, such as the 
new genera Ikedosoma and Maxmulleria. It is to be hoped that 
Dr. Bock will continue his fundamental work and will be able to 
revise the genus Ochetostoma, badly in need of an overhauling. 

As Dr. Bock's scheme of classification differs from mine, it is given 
herewith in skeleton form: 

Class EcHiuHOiDEA [of phylum Annelida] 
I. Order Echiuroinea, nov. 

1. Family Echiuridae Baird, 1868. Genera: Echiurus and Urechis. 

2. Family Thalassematidae, nov. 

a. Subfamily Ikedinae, nov. Genus: Ikeda. 

b. Subfamily Thalassematinae, nov. Genera: Thalassema, Ocheto- 

stoma (incl. Li stri Globus) , Ikedosoma, Arhynchite. 

3. Family Bonelliidae Baird, 1868. Genera: Maxmiilleria, Acantho- 

hamingia, Archibonellia, Hamingia, Parabonellia, Protobonellia, 
Pseudobonellia, Bonellia. 
JI. Order Saccosomatinea, nov. 

1. Family Saccosomatidae Theel, 1906. Genus: Saccosoma. 
III. Order Poeobiinea, nov. 

1. Family Poeobiidae Heath, 1930. Genus: Poeobius. 

"The two latter orders comprise each a single species and they 
must be regarded as very aberrant Annelids of somewhat doubtful 
relationship to the true Echiuroids" (p. 17). 



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ish water with the description of a new species from the Andamans. 
Mem. Asiatic Soc. Bengal, vol. 6, pp. 323-338, 1 pi. (Thalassema 
kempi.) 
1919b. Notes on echiuroids from Chandipore, Orissa. Rec. Indian Mus., 

vol. 16 pp. 399-402. {Thalassema microrhynchus.) 
1920. On a new species of Thalassema from the Gulf of Manaar with notes 
on Thurston's species T. formulosum. Rec. Indian Mus., vol. 19, 
pt. 2, pp. 35-37. (Thalassema hornelli.) 
1935. On a collection of echiuroids of the genus Thalassema Lamarck in the 
Indian Museum, Calcutta. Rec. Indian Mus., vol. 37, pt. 1, pp. 
39-43, 1 pi. {Thalassema arkati, T. marshalli.) 
Prashad, B., and Awati, P. R. 

1929, On a new species of the genus Thalassema from Bombay. Rec. In- 
dian Mus., vol. 31, pt. 4, pp. 259-261, 1 pi. (Thalassema bombay- 
ensis.) 

QUATREFAGES, JeaN LoUIS ArMAND DE 

1865. Histoire naturelle des annells marins et d'eau douce, vol. 2, pp. 591- 

599, pi. 16, fig. 13. 
Redfield, Alfred C, and Florkin, Marcel. 

1931. The respiratory function of the blood of Urechis caupo. Biol. Bull., 

vol. 61, pp. 185-210, 5 figs. 

RiETSCH, MaXIMILIEN. 

1886. fitude sur les g6phyriens arm^s ou ^chiuriens. Recueil Zool. Suisse, 
vol. 3, No. 3, pp. 313-515, 6 pis. 
RiCKETTs, Edward F., and Calvin, Jack. 

1939. Between Pacific tides, 320 pp., 112 figs., 46 pis. Stanford University, 
Calif. 
Sato, Hatao. 

1931. Report of the biological survey of Mutsu Bay. 20, Echiuroidea. Sci. 
Rep. Tohoku Imp. Univ., ser. 4, vol. 6, pp. 171-184, 4 figs. 

1934. Report on the Sipunculoidea, Echiuroidea and Priapuloidea collected 

by the Soyo-Maru Expedition of 1922-1930. Sci. Rep. Tohoku 
Imp. Univ., ser. 4, vol. 9, pp. 1-32, 31 figs., 1 pi. 

1935. Sipunculoidea and Echiuroidea of the West Caroline Islands. Sci. 

Rep. Tohoku Imp. Univ., ser. 4, vol. 10, pp. 299-329, 17 figs., 3 pis. 



ECHIUROID WORMS OF NORTH PACIFIC — FISHER 285 

Sato, Hayao — Continued 

1937. Echiuroidea, Sipunculoidea and Priapuloidea obtained in northeast 
Honshu, Japan. Saito Ho-on Kai Mus. Res. Bull., No. 12, pp. 
137- 17G, 3 pis. 
1939. Studies on the Echiuroidea, Sipunculoidea and Priapuloidea of Japan. 
Sci. Rep. Tohoku Imp. Univ., ser. 4, vol. 14, pp. 339-400, 60 figs., 
5 pis. 
Seitz, Philipp. 

1907. Der Hau von Echiunis chilcnsis {Vrechis n. g., chilensis). Zool. 
Jahrb., Abt. Anat., vol. 24, pp. 323-350, 3 pis. 
Selenka, Euil. 

1885. Report on the Gephyrea collected during the voyage of H. M. S. 
Challenger. Challenger Reports, vol. 13, pp. 1-24, 4 pis. 
Shipley, Arthur. 

1899. On a collection of echiuroids from the Loyalty Islands. Willey's 

Zoological Results, pt. 3, pp. 335-356, 1 pi. (Review of nominal 
species of Echiuroidea.) 
Skorikov, a. S. 

1909. Subfam. Echiurini nov. (Gephyrea armata). Ann. Mus. Zool. Acad. 
Imp. Sci. St.-P6tersbourg, vol. 14, pp. 77- 102, 1 pi. 
Sluiter, G. p. 

1883. t)ber einige Sternwlirmer des indischen Archipels. Zool. Anz., vol. 6, 
pp. 222- 228. 
Spengel, Johann Wilhelm. 

1879a. tlber die Organisation des Echiurus pallasii, I. Zool. Anz., vol. 2, 

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1879b. Beitrage zur Kenntniss der Gephyreen, I: Die Eibildung, die Entwick- 
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vol. 1, pp. 358-419, 5pla. 
1880. Idem, II: Die Organisation des Echiurus pallasii. Zeitschr. wiss. 

Zool., vol. 34, pp. 458-538, 4 pis. 
1912a. Idem, III: Zum Bau des Kopflappens der armaten Gephyreen. 

Zeitschr. wiss. Zool., vol. 101, pp. 342-385, 21 pis. 
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1912c. t)ber den Ilautmuskelchlauch gewisser Thalassema Arten und Seine 
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GeH., vol. 22, PI). 309-317. 
Steinbeck, John, and Ricketts, Edward F. 

1941. The s*;a of Cortoz, 598 i)p., 40 pis. New York. 
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1906. Northern and Arctic invertebrates in the collection of the Swedish 
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Wharto.v, L. D. 

1913. A description of sorno Philippine; Thalassemac with a revision of the 
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1900. Our North American echiurids. Biol. Bull., vol. 1, pj). I63-17H, 1 pi. 



EXPLANATION OF PLATES 

All figures were made by the writer directly from dissections or 
specimens. 

Plate 20 

Echiurus echiurus alaskanus, new subspecies 

1, Dorsal view of anatomy of anterior portion of body, X 5. Body wall is indi- 

cated only in pharyngeal region. Junction of diaphragm to body wall is 
indicated by dashes for dorsal half and dots for ventral half. Coiled loops 
of esophagus are shown emerging from posterior opening of diaphragm, 
below the interbasal muscle. Lines radiating from pharynx are the muscu- 
lar frenula. The peripharyngeal coelom is shaded. 

2, Anal bristle from posterior ring, 7.5 mm. long. 

3, Anal bristle from anterior ring, 8 mm. long. 

4, Diagram of the diaphragm (ventral half dotted) showing the posterodorsal 

opening. The esophagus is omitted except where it pierces the right wall 
and becomes the gizzard. The edges that are attached to body wall and 
ventral mesenteries are dashes. 

5, Diagram of a section through diaphragm showing how its lower border merges 

with the ventral mesenteries. 

B^B*, dorsal, ring, neurointestinal, and ventral blood vessels, respectively," 
C, stomach; CG, ciliated groove; D, diaphragm; G, gizzard; MD, dorsa 
mesenteries of esophagus; MV, ventral mesenteries of esophagus (fig. 5); 
A^, nephridium; NC, nerve cord; 0, esophagus, indicated by arrows, ventral 
mesentery omitted, dorsal inesenteries shown as lighter lines; P, pharynx; 
Si, anterior end of siphon; X, perivisceral coelom (Sg. 4); A^^, peripharyngeal 
coelom. 

Plate 21 

Listriolohus pelodes, new species 

1, Ventral view of a large specimen from Monterey Bay, Calif., X l}4. 

2, Same specimen, X 5, ventral view of anterior end showing nerve loop in pro- 

boscis, a section of which has been removed. 

4, Small phase before muscle bands are evident, natural size. 

4c, Side view of anterior end of a living specimen, X 3. 

46, A female, X 3, showing nephridia and fecal pellets. At this size the muscle 
bands are not apparent unless the specimen is stronglj^ contracted. Out- 
line from living animal. 

Ocheiostoma octomyotum, new species 

3, Ventral view of type specimen from Newport Bay, Calif., natural size. A 

specimen from Cabrillo Beach, near San Pedro, 95 mm. long, has a pro- 
boscis 93 mm. 

Plate 22 

Listriolohus pelodes, new species 

Type specimen, X 7, dissected to show organs of anterior portion of the body; 

the alimentary canal is drawn to the right of its natural position. 
B', dorsal blood vessel; B^, ring vessel; B^, neurointestinal connective; B*, ventral 
vessel; C, stomach; CG, ciliated groove of intestine; G, gizzard; I, intestine; 
286 



ECHIUROID WORMS OF NORTH PACIFIC — FISHER 287 

Ml, iiitorbftsal muscle of sctao; X, iiophridiiiin; \C, nerve cord; 0, esoph- 
ag:iis; P, pharynx; P\ posterior end of pharynx; 8, seta;«Si/, anterior end of 
siphon; VM, ventral mesentery; X, parasite. 

Plate 23 

Ochetostoma octomyotum, new species 

1, Dissection, X 2; the spiral funnels of the nephridia and longitudinal muscle 

hands have been omitted except the midventral, which is lighter shaded. 

2, Interval between right ventrolateral and lateral muscles at middle of body 

showing fascicles of the oblique layer. On the lower left corner the oblique 
layer has been removed, X 20. 

3, Cloaca and adjacent part of intestine opened to show relation with intestinal 

coecum and anal vesicles, X 5. 
Lettering as for plate 24. 

Pl.\te 24 

Ochetostoma octomyotum, new species 

The anterior portion of plate 23, figure 1, enlarged X 5 and with addition of 
details. 

A, anus; AV, anal vesicles; AV\ their opening into cloaca; B', dorsal blood ves- 
sel; D*, ring vessel; IP, neurointestinal connective of which B^ is merely a 
part; B*, ventral vessel; C, stomach; CF, ciliated funnel or nephrostome; 
CG, ciliated groove of intestine; CI, cloaca; DM, dorsal mesentery of pharynx, 
G, gizzard; IC, intestinal coecum; IC\ its opening into cloaca; MC, outer 
circular muscle layer; MD, dorsal muscle band; MDL, dorsolateral muscle 
band; ML, lateral muscle band; MO, oblique inner layer of muscles; MVL, 
ventrolateral muscle band; N, nephridium; NC, nerve cord; O, esophagus; 
P, pharynx; S, seta; Si, siphon; Sil, entrance to siphon; Si2, end of siphon; 
VM, ventral mesentery of pharynx. 

Plate 25 

Arhynchite inamoenua, new species 

1, \'entral view of paratype, X 1. 

2, Seta of type, X 10. 

3, Di.s.section of anterior complex of type, X 7, showing foregut in situ. 

4, Skin of figure 1, from near midventral lino, enlarged. 

."J, TyjK>, X 7; the interbasal muscle has been cut aii<l the liberatod foregut drawn 

to the right; ventral mesentery is dotted. 
B', B*, B*, dorsal, neurointestinal, and ventral blood ves.sels, respectively; CF, 

nephrostome; C, gizzard; M, mouth; MI, interbasal muscle; A^, nephridium; 

NC, nerve cord; O, e.soi>hagus; P, ])liarynx; S, 8(;ta. 

Plate 2G 

BonelliopKig aUiHknnn, new geriu.M and sjjecies 

1, Di.s.Hoction of si)ecirnen from I'liala-Hka, dorsal view, X 4. 

2, Ti]) of one of the priinar\- l)rancheH of an anal vesicle, showing two secondary 

branches with their ciliated funnels, X 50. 

3, One of the ciliated funnels, X 200. 

4, Nephrostome, X 10. 



288 PROCEEDINGS OF THE NATIONAL MUSEUM vol. se 

5, Anterior end of an individual, the short proboscis apparently in process of 

regeneration, X 3. 

6, Ventral view of a worm life size; the second proboscis indicates about the 

maximum length in preserved specimens. 

7, Anterior ventral portion of worm showing the contracted verrucose skin, X 10; 

when the skin is fully distended the verrucae disappear, the glandular thick- 
enings then appearing as squarish patches, closely spaced. 
Go, gonad; other lettering as for plate 24. 

Plate 27 

Bonelliopsis alaskana, new genus and species 

1, Dissection of anterior end of a specimen with a right nephridium filled with 

eggs, X 5. A "window" has been cut in the dorsal wall of pharynx and also 
one in the nephridium to show the invaginated tip, within which is the 
nephrostome. The longitudinal ridges on inner wall of nephridium are indi- 
cated by dotted lines. 

2, Same, anterior end, in contracted state, with right half removed to show 

relation of dorsal and ventral mesenteries to alimentary canal and ventral 
mesenteries to alimentary canal and nephridium, X 5. Anteriorly only a 
few of the frenula of pharynx indicated; back of these the transverse mesen- 
tery of figure 1 (M' ) is indicated as a black line. DM, dorsal mesentery, and 
DM^ (dot-dash), its attachment to right wall of body (removed); VM, 
ventral mesentery, attached to lower side of intestine and mostly to left side 
of f oregut and involving posterior part of dorsal artery (B') . The nephridium 
lies in a sort of anterior cul-de-sac or egg trap. When the animal is extended 
the upper margin of dorsal mesentery is at a distance behind the nephridium, 
here shown at minimum size. 

Af' , a transverse mesentery below the dorsal blood vessel; 

M', special muscular mesentery between gizzard and stomach, not shown in figure 
2; PV, position of pyloric valve, mentioned in text; other lettering as for 
plate 24. 

Plate 28 

Eubonellia valida, new genus and species 

1, Dissection of anterior end of body from above, X 4. The f oregut has been 

drawn to the left to show the large nephridium (N) with its terminal nephro- 
stome (CF). Note the unusually long stomach (C), filled with pellets, 
between the gizzard (G) and intestine (/). The ventral blood vessel (B*) 
and the much contracted neurointestinal vessel (B') have been cross- 
hatched. The dorsal blood vessel (B') is unshaded. 

2, A cluster of nephric elements of an anal vesicle, X 30. 

3, Type, natural size, from below. 

4, A pellet from the postsiphonal intestine, X 10. 

5, Male, from mouth cavity, X 50; anterior end to right; the sperm receptacle and 

duct shown. 
Lettering as for plate 24. 

Plate 29 

Nellobia eusoma, new genus and species 

1, Ventral view of anterior end of type showing the short truncate^snout, X 5. 

2, Ventral view of type, X 1. 



ECHIUROID WORMS OF NORTH PACIFIC — FISHER 289 

3, Teruiinal portion of intestine, the cloaca, anus, and anal vesicles, X 5. 

4, Branchlot of anal vesicle, X 50. 

NP, nephridiopore; Go, posterior part of gonad with blood vessel and nerve cord 
underneath; M, mesenteries; other lettering as for plate 24. 

Plate 30 

Nellobia euaoma, new genus and species 

1, Dissection of anterior portion of type, seen from above, X 5. The nephridium, 

filled with eggs (0.85 mm. in diameter) on the left, has a window cut in the 
wall to show the constricted duct from the egg chamber. The swollen duct 
leading to external opening lies under the nerve cord and ventral blood 
vessel. 

2, Pharynx, X 5. Interior; anterior end looking toward mouth. The dorsal side 

has been cut open. 
B\ B*, B*, dorsal, neurointestinal, and ventral blood vessels, respectively; CF, 
nephrostome; D, peripheral portion of peripharyngeal diaphragm (the central 
portion adjacent to pharynx has been removed); DAI, dorsal mesenteries; 
G, gizzard; Go, gonad; M, mesenterial sheet holding loop of pharynx-esopha- 
gus; NC, nerve cord; O, esophagus; Sil, beginning of siphon; VM, ventral 
mesentery of pharynx-esophagus; X-Y, probable extent of gizzard; at Y 
the canal was broken, and it is possible that a portion of the succeeding 
stomach was lost. 

Plate 31 

Acanthohamingia paradola, new species 

1, Ventral aspect of paratype, X 1. 

2, Same specimen; genital groove, extending forward from nci)hridiopore (AO 

and showing four males in situ, X 5 ( $ , males attached to skin). 
2a, Anterior portion of figure 2, X 15. 

3, Type; genital groove in probably the normal closed state, X 5. 

4, Genital groove of third specimen that has two nephridiopores {N), X 5. 
4a, Nephridiopores of above, enlarged. 

5, Male from genital groove of paratype (fig. 2), 1.19 mm. long, X 50. 

6, Tyj)e; anal vesicles and thin-walled cloaca from above, X 3. The anus can be 

seen through the thin wall of the cloaca, and on the left most of tlie tubes 
spring from a rudimentary bladder. 
I, intestine Cmi.ssing from type); M, mouth; A'^, nephridiopore; S, spermotheca 
(nephridium). 

Plate 32 

Acanthohamingia paradola, new species 

Dissection of anterior part of animal from above, X 4. Note the very long forogut 
ending at B^ and the long (as compared with Bonellia) segment of intestine 
between B^ and Sil. 

B\ B*, B*, dorsal, neurointestinal, and ventral blood vcssel.s, respectively; C, 
portion corresponding to stornarli of other bonelliids; CF, nephrostome; G, 
probable gizzard; /, intestine; A^, ncphridia; NC, nerve cord; 0, esophagus; 
P, pharynx, Pe, pellet, X 10; Si, siphon; Sit, anterior end of siphon. 



290 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

Plate 33 

Urechis caupo Fisher and MacGinitie 

1, Dissection of contracted specimen from above, showing the intestine in hap- 

hazard convolutions. The principal mesenterial bands, which anchor the 
intestines to the body wall, are shown but not lettered. The coelomic aper- 
tures of the nephridia are recognizable by the conspicuous coiled lips. The 
arrow indicates point where pharynx becomes esophagus. The figures in 
sequence on the midgut are intended to aid in following the convolutions; 
1 is at the beginning and IS near the end. Beyond 13 the slight diverticulum 
of the respiratory gut is indicated. 

2, Pharynx contracted and slit open along ventral side to show the straight 

longitudinal folds of lining, continuous with those of the proboscis. Pos- 
teriorly is shown characteristic lining of esophagus; on each side are the 
dorsal mesenteries. In front of these the ventral mesenteries are spread 
laterally since pharynx has been opened ventrally (see pi. 34, fig. 2, 14)- 

3, Ventral surface of proboscis and anterior end of body. 

4, Posterior end of body showing eccentric anus and circle of setae ; + marks the 

midventral line. 

5, Two views of the anterior setae. The line indicates 1 mm. 

6, An anal seta, same scale as figure 5, with, below, a tip enlarged. 

AS, anterior setae (accessory seta shown at side; muscles not drawn); AV, anal 
vesicles; Ci, C2, crops 1 and 2 of Seitz, subtended by muscular bands Mi and 
M2; C2 is the stomach; CL, cloaca, the posterior portion lined with heavy 
longitudinal ridges (arrows mark apertures of anal vesicles) ; G, gizzard, a 
portion of the foregut lying between Ci and C2, characterized by thick 
muscular walls and circular muscular ridges and constrictions; HG, respira- 
tory gut, specialized posterior segment of midgut; Mi, M2, muscular bands 
of crop and stomach (C2) ; A'', nephridium; NC, nerve cord; 0, esophagus, 
anterior limit marked by an arrow; P, pharynx; PS, posterior or anal setae; 
S, siphon or accessory intestine; SI, beginning of siphon near beginning of 
midgut; S2, end of siphon; VM, ventral mesenteries of pharynx; 1-13, these 
figures are in sequence along the midgut and are intended to aid in following 
the course; 13 is near the junction of midgut and its terminal specialized 
portion, the respiratory gut. 

Plate 34 

Urechis caupo Fisher and MacGinitie 

1, Dissection of contracted individual showing the generically characteristic 

parts of alimentary canal, most of the "small intestine" having been removed. 
The very long foregut consists of pharynx (anterior to P), esophagus (0), 
crop (C) with its strong muscular mesentery, gizzard (G), and stomach 
(C^) anchored posteriorly by a strong mesentery here shown in maximum 
contraction. Si is beginning of siphon (pi. 35, fig. 5). Attachment of 
respiratory gut is always on left of nerve cord (NC) . Along its dorsal surface 
_ is shown the muscle strand continued from the small intestines and serving 

posteriorly for attachment of a few dorsolateral mesenteries. 

2, Head region of coelom looking forward from just behind first pair of nephridia 

{16) showing arrangement of muscular mesenteries of pharynx: 1-7, dorsal 
and dorsolateral; 8-12, lateral and ventrolateral; 13, the dorsoventral 
muscles mentioned in text; I4, ventral mesenteries of pharynx; 17, nerve 



ECHIUROID WOKMS OF NOUTH PACIFIC — FISHER 291 

cord; crossiiiu; the ventral mesenteries between the two figure i.{'s is the 
interbasal muscle of setae; radiating muscles of setae shown in solid black 
on right. 

3, Anterior ainpoct of a nephridium of second pair showing nephrostome and 

elongated lips spirally coiled, X 5. 

4, A nephridium from a specimen 40 mm. long, not yet sexually mature, X 20. 

Plate 35 
Urechis caupo Fisher and MacGinitie 

1, Arrangement of intestine and mesenteries in fully expanded sjx'cimen, X ^. 

Anterior and posterior portions of body have been omitted and the intestine 
has been spread to right and left to show attachments. Normally these 
lateralmost parts overlie the darker and more mesially located portions. A 
section has been removed from respiratory gut to show attachment of mes- 
enteries. C, crop; C^ stomach; G, gizzard; IIG, respiratory gut; M^, 
mesentery of stomach; NC, nerve cord; Si, siphon; Sil and Si2, anterior and 
posterior end of siphon. 

2, Portion of midgut at A' of figure 1, showing its highly sacculate structure; 

mucosa with fine anastomosing plications, transverse in direction; Si, 
siphon, X 2. 

3, Postsiphonal midgut at XX of figure 1, X 2, showing the mucosa and longitudi- 

nal muscle band marking position of ciliated groove. 

4, 4a, Cestode larvae from anterior end of the siphon, X 60. These larvae vary 

in length from 0.25 to 0.32 mm. and are free in the lumen of siphon and in 
the hernialike swellings, which may be caused by them (.see fig. 5, above Sil). 

5, Sagittal section, X 5, of the distal end of stomach and beginning of midgut and 

siphon, showing macroscopic character of mucosa; C^, stomach; CG, one side 
only the ciliated groove; the groove is constituted by two of these finely 
plicated folds or ridges of the mucosa. The plications are coarser and the 
groove is broader in the short segment CG'; Si, siphon, showing foliose 
mucosa. The cestode larvae were found in this portion and in the hernia- 
like swellings shown just above SjI, the narrow passage connecting midgut 
and siphon. 

6, Fecal pellets, X 3. Specinjen from Monterey Bay, Calif. 

Plate 36 

1, I'rrchin chilcnaia (Miillcr): Anterior end of respiratory gut showing mode of 

attachment by muscular mesenteries, X 3. Compare wit It figures 2 and 3. 

2, r'rec/ii« c«i/po Fisher and MacGinitie, X 1.5. 

3, IJrechis unicinctu8 (von Drasche), X 5. 

4, L'rechin caupo: Cloacal region of a relaxed specimen (pi. 35, fig. 1) with the 

posterior part of respiratory gut ; dorsal wall of cloaca removed. ..1 V, anal 
vcHicle; M^, nmscular masentery anchoring the stomach; M*, muscular 
fronula of cloaca; A^, ncphridia; NC, ncrvo cord; PS, posterior setae. 

Plate 37 

Urechis caupo anfl Conmiensals 

1, Portion of tunnel showing one position of worm while pumping water through 
its slime-net and characteristic stations of commen.sals, X *i. A, Cleve- 
landia ion (Jordan and Gilbert) at mouth of tube; B, IlespcronoS advrntor 
(Skogsberg) ; C, Scleroplax granulata Rathbun; D, Cnjplomya californica 
(Conrad). At U|)i)er [)f)irit where tube is interrupted one inch has been 
omitted, at lower point four inches. 



292 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 8« 

2, Scleroplax granulata Rathbun, male, X 3. 

3, Hesperonoe adventor (Skogsberg), type X 1. 

4, Specimen of U. caupo with slime-tube in place, with the thickening at point of 

attachment to body indicated. The worm is shown in characteristic pump- 
ing posture; X 3. 

5, Characteristic posture while tube is being secreted; tube just begun; entrance 

indicated by dots; X 1/4. 

6, Expression of worm while swallowing slime-tube. 

7, Grasping slime-tube at moment of starting to swallow. The proboscis is 

holding the posterior edge of tube while a portion is being sucked in on 
ventral side. This step occupies about three seconds. 



U. S. BOVERHMENT rRINTIN* OFFlCli l»4« 



S NATIONAL MUSEUM 



PROCEEDINGS VOL "^6 PLATE 20 




LCMIURUS ECHIURUS ALASKANUS. NEW ^UOsHH.Itt. 
POn t.Xfn.*NATION SEE PAOE ?«« 



U. S. NATIONAL MUSEUM 



PROCEEDINGS, VOL. 96 PLATE 21 




LISTRIOLOBUS PELODES. NEW SPECIES. AND OCHETOSTOMA OCTOMYOTUM, NEV 

Species 

FOR EXPLANATION SEE PAGE 286 



U. S. NATIONAL. MUSEUM 



PROCEEDINGS. VOL 96 PLATE ?2 

< 




LISTRIOLOBUS PELODES. NEW SPECIES 
roB i;kpi_anation see paocs a»» m 



U. S. NATIONAL MUSEUM 



PROCEEDINGS, VOL. 96 PLATE 23 




OCHETOSTOMA OCTOMYOTUM, NEW SPECIES 

FOR EXPLANATION SEE PAGE 287 



U S NATIONAL MUSEUM 



PROCEEDINGS. VOL 96 PLATE 24 




MV)7-*^\J MVL 



^ ML ^M 



DL 



OCHETOSTOMA OCTOMYQTUM Nf W ijPtCIES 
row tXPLANATION SEE PAGE .'»? 



U. S. NATIONAL MUSEUM 



PROCEEDINGS. VOL. 96 PLATE 25 




ARHYNCHITE INAMOENUS, NEW SPECIES 
FOR EXPLANATION SEE PAGE 287 



U S NATIONAL MUSEUM 



PRCX:EEDINGS. vol. 96 PLATE 26 







BorJLLLIOPSIS ALASKANA. NEW GENUS AND SPECIES 
FOn FXPUANATION SEE PAr.l 



U. S. NATIONAL MUSEUM 



PROCEEDINGS, VOL. 96 PLATE 27 




BONELLIOPSIS ALASKANA. NEW GENUS AND SPECIES 
FOR EXPLANATION SEE PAGE 288 



U S NATIONAL MUSEUM 



PROCEEDINGS. VOL. 96 PLATE 28 




EUBONELLIA VALIDA. NEW GENUS AND SPECIES 
FOR EXPLANATION BEE PAGE /»» 



U. S. NATIONAL MUSEUM 



PROCEEDINGS. VOL. 96 PLATE 29 



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Nellobia eusoma. New Genus and species 

FOR EXPLANATION SEE PAGES 298-289 



U S. NATIONAL MUSEUM 



PROCEEDINGS. VOL. 96 PLATE 30 




DM 



f-JhLioBiA tijsoMA. New Genus and Species 

FOR EXPUANATION Str PACiE I<W 



U. S. NATIONAL MUSEUM 



PROCEEDINGS, VOL. 96 PLATE 31 



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ACANTHOHAMINGIA PARADOLA, NEW SPECIES 

FOR EXPLANATION SEE PAGE 289 



U S NATIONAL MUSEUM 

r 



PROCEEDINGS. VOL 96 PLATE 32 




ACAN f HtJHAMINICIA PARADOLA NEW SPECIES 

ro« FxriANATiOfj sie I'A'.c ;;•» 



U. S. NATIONAL MUSEUM 



PROCEEDINGS. VOL. 96 PLATE 33 







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F^<:4- 



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Urechis caupo Fisher and MacGinitie 

FOR EXPLANATION SEE PAGE 290 



U S NATIONAL MUSEUM 



PROCEEDINGS. VOL S6 PLATE 34 




URECHis CAUPO Fisher and MacGinitie 

rOR eXPl^NATION r.EE PACES .■»- »1 



U^ S. NATIONAL MUSEUM 



PROCEEDINGS. VOL. 96 PLATE 35 




Urechis caupo Fisher and MacGinitie 

FOR EXPLANATION SEE PAGE 291 



U. S NATIONAL MUSEUM 



PROCEEDINGS VOL 96 PLATE 36 




unlchis caupo. U chilensis. and U. UNICINCTUS 

FOR EXPLANATION SCC PAGB 291 



U. S. NATIONAL MUSEUM 



PROCEEDINGS. VOL. 96 PLATE 37 




URECHIS CAUPO AND COMMENSALS 
FOR EXPLANAXrON SEE PAGES 291-292 




PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 



by the 



SMITHSONIAN INSTITUTION 
U. S. NATIONAL MUSEUM 

Vol. % W.>hington: 1946 No. 3199 



THE OSTEOLOGY OF THE FOSSIL TURTLE TESTUDO 
PKAEEXTANS LAMBE, WITH NOTES ON OTHER 
SPECIES OF TESTUDO FROM THE OLIGOCENE OF 
WYOMING 



By Charles W. Gilmore * 



Among the fossil vertebrate materials collected from the Oligocene 
of the Indian Creek Basin area in Wyoming by the Smithsonian 
Paleontological Expeditions of 1932 and 1942 were six well-preserved 
specimens of the land tortoise Testudo. Interest in these specimens 
centei-s primarily in the fact that they contribute the first information 
to be had of the skull and internal skeleton of Testudo j^raeextans 
Lambe, as well as illustrating tlie individual variation found in the 
shell structure of that species. Two of these specimens pertain to 
Testudo laticunea Cope, this being the first time this species has been 
recognized in this area. 

Family TESTUDINIDAE 

Genus TESTUDO Linnaeus 

TESTUDO PKAEEXTANS Lambe 

Plates 38-41; Platk 44, Fkjckk 2 

Testudo praeextana Immbk, Ottawa Nat., vol. 27, j)!). '>7 (W, |)ls. 4, !">, 1913. — Hay, 
Carnegie Inst. Washington Pul.i. 390, p. 101, 1930. 

Diagnosis. — The cliaractcrs that at tliis time appear to (iistiii^'uisii 
Testudo praeextans arc a.s follows: Cara{)ace depressed, with Jhitteiied 
toj) in the vertebral region; prominent epiplastnd lip, project iiig 
beyond the borders of the curai)ace and with j)arallel sides; sulci 
usually shallowly impressed and usually ending, on free borders, in 

'.Mr. fJilmorc died on 8vptomh«r 27, 104fi.— Ki>. 

873370— IC 1 -'03 



294 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

a projecting point or miicro; neurals variable but usually less dif- 
ferentiated than in most species of the genus; first neural longest 
of the series; entoplastron pointed in front, rounded behind; gular 
scutes constantly encroaching on the entoplastron ; suprapygal usually 
in contact with the eleventh peripheral only, rarely reaching the 
pygal; median longitudinal ridge in front of the choanae. 




Figure 20. — Skull of Testudo praeextans Lambe (U.S.N.M. No. 15874), superior view: 
exoc, exoccipital; /, frontal; /, jugal; mx, maxillary; occ, occipital condyle; op, opisthotic; 
p, parietal; pf, prefrontal; pmx, premaxillary; prf, postfrontal; so, siipraoccipital; sq, 
squamosal. Natural size. 

Materials. — ^The four specimens pertaining to Testudo praeextans 
are all from Niobrara County, Wyo., and from the Brule division of the 
Oligocene. The best-preserved specimen, U.S.N.M. No, 15874, col- 
lected in 1932, consists of the complete carapace, plastron, skull, 
lower jaws, pectoral and pelvic girdles, both humeri, both ulnae, one 
radius, both femora, both tibiae, one fibula, and parts of all four feet. 
It was found on the Anderson Ranch on the south side of Young 
Woman Creek; specimen U.S.N.M. No. 15878, collected in 1932, was 
found on the east side of Little Indian Creek and consists of the com- 
plete shell, parts of both humeri, both ulnae, both radii, one articulated 
forefoot, and portions of the other three forefeet; specimen U.S.N.M. 



OSTEOLOGY OF TESTUDO PRAEEXTANS — GILMORE 



295 



No. ir»72S was foiiiul in 1942 about a inik" north of Wliitnian Post- 
oflice and consists of carapace, plastron, hunuMus, both scapulae, both 
coracoids, both tii)iao, both fibulao, tarsals, and sonic foot bones; 
specimen I'.S.X.M. Xo. 10)732 was coUccti'tl in 1942 on the Anderson 
Kanch on the nortli side of Young Woman Creek and consists of 
carapace, })lastron. pelvis, two humeri, coracoid, two scapulae, tibia, 
two ulnae, two radii, parts of both fibulae, and foot bones. 

DESCRIPTION 

Skull. — Of the 48 or more species of Testudo described from North 
America, the skulls of only sLx are known at the present time. These 
are Testudo thom.soni Hay, T. peragrans Hay, T. oshorniana Hay, T. 
impensa Hay, T. orthopygia (Cope), and T. gilberti Hay; and only one 
of these, T. thomsoni, is from the Oligocene. To this list we may now 
add T. jiraeertans Lambe, represented by an unusually well preserved 
skull and lower jaws belonging to specimen U.S.N.M. No. 15874 and 
illustrated in three views in figures 20-22. 




Figure 21. — Lateral view of skull and lower jaws of Testudo praeextans Lanibc (U.S.N.M. 
No. 15874): an, angular; ar, articular; c, coracoid; d, dentary; /, frontal; /, jugai; mx^ 
maxilla; occ, occipital condyle; p, parietal; pf, prefrontal; p7nx, premaxillary; prj, post- 
frontal; qu, quadrate; sa, surangular; so, supraoccipital; sq, squamosal. Natural size. 

The skull and lower jaws of this specinicn arc practically complete 
and undistorted, with numy of the cranial sutures clearly distinguish- 
able (fig. 21). Th(! skull is of moderate size. Its length, from the tip 
of the premaxillary to the occipital condyle, is 81 mm.; its width across 
the sf|UHinosals is 57 mm.; the height from tiie cutting edge of the 
mu.xiila to the upper surface of the frontal is 2\\ mm. 'V\\v. sides of the 
skull forward to the backs of the orbits arc ncniiy parallel; beyond 
here tliev regtdarly converge to the broadly rounded snout. Viewed 
from the side tiie upper outline of the skull is nearly straight to a i)oint 
above the anterior borders of the orbits where the prefrontals bend 
strongly downward. 



296 



PROCEEDINGS OF THE NATIONAL MUSEUM 



The prefrontals meet along the median line, a distance of 16 mm., 
and this is also the length of the suture between the frontals. The 
lateral angles of the skull extend backward to a point slightly posterior 
to the occipital condyle. The orbit has a greatest anteroposterior 
diameter of 23 mm.; the nasal opening is 17 mm. wide; the interorbital 
space is 22 mm. wide. 

Viewed from above, the skull of Testudo praeextans has its closest 
resemblances in the cranium of the Miocene T. osborniana, differing 




Figure 22. — Palatal view of the skull of Testudo praeextans Lambe (U.S.N.M. No. 15874): 
bo, basioccipital; bsp, basisphenoid; ect, ectopterygoid; mx, maxillary; occ, occipital 
condyle; pi, palatine; pmx, premaxillary; pt, pterygoid; qu, quadrate; so, supraoccipital; 
sq, squamosal; v, vomer. Natural size. 

in its smaller size, the more abruptly truncated snout, the more 
bluntly pointed squamosal region, the longer supraocciptal process, 
and the straighter cutting edge of the maxillary. The orbit also has 
a greater anteroposterior diameter, being one-fourth the over-all 
length of the skull. The anteroposterior extent of the otic region 
measured across the paraoccipital and the prootic is 22 mm. 

The incomplete skull of Testudo thomsoni, the only other Oligocene 
species in which the cranium is now known, differs from the skull 



OSTEOLOGY OF TESTIDO PHAKKXTANS — GILMORE 297 

before me in hav^ing shorter prefontals, smaller frontals, and longi- 
tudinal channels on the masticatory surfaces unequal in width. 
Wlii'ihcr those diU'eronces constitute stable characters that can be 
relied upon for their specific separation cannot be determined until 
more skulls ai-e available. 

The roof of the mouth is moderately vaulted and traversed along 
the midline by a sharp ridge (fig. 22), the anterior end of which abrupt- 
ly widens where the vomer articulates with tiie premaxillaries and 




SQ 



Figure 23. — Lower jaw of Testudo praeextans Lainbe (U.S.N.M. No. 15874), superior view: 
ar, articular; c, coronoid; d, dentary; prar, prearticular; sa, surangular. Natural size. 

maxillary bones. Posteriorly this vomerine ridge underlaps the 
pterj-goid part of the ridge, which merges into the palatal surfaces 
of the pterygoid slightly in advance of their median union with the 
basisphenoid. 

The masticatory surface of the ma.xillary is divided by a low metlian 
ridge that is received in a groove of the mandible. This ridge is 
bordered on each side by longitudinal grooves of about equal width. 
In this respect this specimen differs markedly from Testudo thomsoni, 
w liich has a very narrow inner groove and a widened out(^r groove. 
The imier ridge is unusually low, and it meets its fellow of the opposite 
side on the posterior border of the premaxillary. At the midline 
along th(! symphysis of the premaxillae there is a prominent longi- 
tudinal ridge, as in Oopherus and in some living species of Testudo. 
This ridge is absent in the skulls of other American fossil Testudo 
now known. It was the sup[)osed absence of this ridge in front of the 
choanae that led Hay to use it as one of the important characters for 
distinguishing Testudo from Stylemys} 

' Hay. O. ['., Fossil turtUs of .North Amcrlcn. rarncRlc trust. Washington Publ. 75. p. 307, \W». 



298 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

The lower jaw, as in Testudo generally, is massive and heavy. Its 
length from the symphysis to the angle is 59 mm., height at the 
coronoid process 59 mm., height at the symphysis 10.5 mm. The 
masticatory surface is traversed longitudinally by a deep groove. 
Inner cutting edge of the mandible is nearly as high as the outer. 
Inner cutting edge of the rami is separated by a notch at the sym- 
physis; cutting edge not denticulated. The extent of the several 
elements forming the mandible is clearly shown in figures 21 and 23. 




Figure 24. — Carapace of Testudo praeextans Lambe, type (C. N. M. No. 8401): 1, 2, 3, 4, 
5, 6, 7, 8, neurals 1-8, respectively. One-sixth natural size. 

Carapace. — In the general contour of the carapace and plastron, 
but more especially in the depressed character of the shell as a whole, 
all these specimens are in close agreement with the type of the species. 
The carapaces of all are flattened on top in the vertebral region and 
when viewed from the side present a nearly straight profile except 
where they curve downward at the ends. This depression of the cara- 
pace, with the exception of Testudo laticunea Cope, appears to dis- 
tinguish this species from all other North American members of the 
genus. Among the large land tortoises only Testudo grandidieri 
VaUlant, of northern Madagascar, has a somewhat simUar depressed 
and flattened shell. 

The four specimens under discussion show a considerable difference 
in size of the shell, as indicated in table 1. 



OSTEOLOGY OF TESTl'DO PR.\KEXTANS — GILMORE 



299 



T.\BLE 1. — Comparative vieaaurements of carapaces of Testudo praeextans 



Specimen 


Greatest 
length 


Oreafest 
width 


Greatest 
height 


Sex 


Width to 
length 


C.N.M. No. 8401 (type).-- 

U.S.N.M. No. 15878- 

r.S.N.M. No. 16728 

r.S.N.M. No. 16732 

r.S.N.M. No. 15874 


Mm. 
479 
485 
525 

' 497 
540 


Mm. 
410 
392 
438 
450 
452 


Mm. 
120 
140 
184 
182 
160 


Female 

Male 

do 

do 

do 


Percent 
85 
80 
83 
90 
83 



• Shortened by crushing. 

Tlic larger size of the National Museum specimens as compaied 
with the type may be accounted for partly by difference in age and 
l)artly by sex. The hollowcd-out plastra of the National Museum 
s])c'cinu'ns clearly indicate them to be males, whereas the flattened 
})lastron of the typo of T. praeextans shows it to be a female. Among 
living land tortoises it is a well-established fact that the males of a 
species attain a larger size than the females. Furthermore, that the 
larger specimens (U.S.N. M. Nos. 15874 and 16732) are aged individ- 
uals is shown by the strong incurving of the pygal region (pi. 40, fig. 
1 ; pi. 41, fig. 2), for it has been observed by Lord Rothschild,^ in his 
study of the Galdpagos tortoises, that "very old individuals of both 
sexes show an inclination often very strong, for the supracaudal to 
curve round towards and even under the posterior end of the 
plastron." 

Viewed from above, all the specimens (pis. 38, 39), including the 
type, are very similar in having their anterior borders broadly rounded 
from side to side; the breadth of the shell decreases more rapidly to- 
ward the front than toward the back, and so the posterior half has a 
squarer outline than the anterior half. This is brought about chiefly 
by the enlargement of the ])eripherals. The posterior border may i)e 
described as angularly lounded, though this contour dilFers c<ui- 
siderably with the age of (Ik; individual. The oldest s])ecimens 
(Nos. 15874 and 10732) are more squarely tniiicate across the pygal 
region tlian the otlieis, owing to the do^viiward and f(tI•^\aI•d deflection 
(»f this ini(l])ortion of the shell. 

The anterior peripherals in all specimens agree in being produced al- 
most horizontally forward; the hiteral ones continue the geiu'ral con- 
vexity of the shell downward, whereas those more posteriorly Mare 
outward and somewhat njjward above the o[)enings for the posterior 
extremities. 

The nuchal, as shown by the measurements in table 2, is fairly con- 
stant in its extent and proportions. The same observation may be 
nuide of the pygal, suprapygal, and second su])rapygal. 

1 liothschlld. I-. W., Nov. Zof.l.. vol. 22. i>. 42N, 1015. 



300 



PROCEEDINGS OF THE NATIONAL MUSEUM 



Table 2. — Comparative measurements {in jnillimeters) of nuchals, pygals, and 
suprapygals of Testudo praeextans 





Nuchal 


Pygal 


Second 
suprapygal 


First 
suprapygal 


Specimen 


Maxi- 
mum 
length 


Maxi- 
mum 
width 


Maxi- 
mum 
length 


Maxi- 
mum 
width 


Maxi- 
mum 
length 


Maxi- 
mum 
width 


Maxi- 
mum 
length 


Maxi- 
mum 
width 


C.N.M. No.8401 (type). 
U.S.N.M. No. 15878_. 
U S N M No. 16728 


95 

82 

102 

107 

102 


103 
103 
113 
113 
113 


70 
66 

74 


62 
76 

82 
84 
83 






84 
81 


102 


51 


74 


99 


U.S.N. M. No. 16732._ 
U.S.N. M. No. 15874_. 


50 
49 


""'74' 


87 
100 


99 

118 



Table 3. — Comparison of shape of neurals of Testudo praeextans 



Neural 
No. 


C.N.M. No.8401 
(type) 


U.S.N.M. 
No. 15878 


U.S.N.M. 
No. 16732 


U.S.N.M. 
No. 15874 


U.S.N.M. 
No. 16728 


1 
2 
3 

4 
5 
6 
7 
8 
9 


Ovate 

Hexagonal 

Quadrangular 
Hexagonal- -. 

do 

do 

do 


Ovate 

Octagonal 

Quadrangular 

Octagonal 

Hexagonal 

do 

do 

do 

do.. 


Ovate 

Octagonal 

Quadrangular 
Hexagonal- -- 

do 

do 

do 

do 


Hexagonal- -- 

do 

Quadrangular 
Hexagonal- -- 

do 

do 

do 

do 


Hexagonal 

Do. 
Quadrangular 
Hexagonal 

Do. 















The free border of the pygal in specimen No. 15874 differs from the 
others in being strongly toothed. The bifurcated first suprapygal 
in the type and in specimen No. 15874 is in contact with the pygal 
and eleventh peripheral, whereas in the other three specimens it 
articulates only with the eleventh peripheral. 

In the shape of the neural bones three features are found common 
to all available specimens: (1) The first neural is the longest of the 
series; (2) the thii'd neural is always quadrangular; (3) the fifth to 
eighth neurals, inclusive, are hexagonal. One individual, No. 15878, 
has two octagonal neurals, the second and fourth; one. No. 16732, 
has only the second neural octagonal; and No. 16728 has none octago- 
nal. In most species of Testudo there are two octagonal neurals, the 
second and fourth, but a series of specimens of other species might 
show them to be equally variable as in T. praeextans. The variation 
in the form of the neurals in T. praeextans is shown in table 3. 

In discussing the neurals of T. laticunea, Hay^ says, "The nem-als 
have not attained so high a degree of differentiation of form as they 
have in most of the species of the genus." If reference is made to the 
absence of octagonal neurals the statement is true of the types of 



3 Hay, O. P., Carnegie Inst. Washington Publ. 75, p. 403, 1908. 



OSTEOLOGY OF TESTUDO PRAEEXTANS— GILMORE 301 

both T. laticunea and T. -praeextans. In my judgment the dirt'orence 
found in the neural bones of the specimens under consich-ration repre- 
sents individual variations well within the species. 

Specimen U.S.N.M. No. 15878 (pi. 38, fig. 2) has an extra neural, 
or nine in all. It is quite evident from an examination of the speci- 
men that the extra element has ileveloped between number 7 and the 
last, which is without corresponding costals. Lambe * has described 
a greater deviation of carapace structure in a specimen of Stylemys 
nebrascensis, which has not only an extra neural but also a ninth pair 
of costal bones and an additional vertebral scute. Tliis same authority 
points out that the type of Stylemys culhertsoni likewise has an addi- 
tional neural. 

The variation in shape of the neurals has brought about a corre- 
sponding variation in the proportions of these bones, as shown in table 
4. In Tf.stvdo praeertans the first neural is consistently the longest of 
any of the series. 

The costal plates in all these specimens are highly modified, and as 
usual in Testudo the second, fourth, and sixth are widened distally and 
narrowed proximally, while the third and fifth are narrowed distally 
and widened at the proximal ends. 

The sulci in most of the specimens are narrow and shallowly im- 
pressed, and where they reach the free edges of the peripherals there is 
usually a small obtuse projection or mucro. 

In the type of Testudo praeextans the sulcus forming the posterior 
boundary of the third vertebral curves strongly forward at the center 
where it crosses the fifth neural, as in U.S.N.M. Nos. 15878, 16728, and 
1G7.32. In specimen No. 15874, however, the sulcus crosses the sixth 
neural as in the type of T. laticunae Cope. 

The proportions of the vertebrals are variable, as shown in tal)le 5. 
In the type of 7'. laticunea and No. 15878 the fifth vertebral is the 
longest of the series, whereas in No. 15874 the first is the longest. 
In the type of T. laticvnea and No. 15874 the first vertebral is longer 
than wide, but in the type of T. praeexiuns and No. 15878 this same 
vertebral is wider than long. No. 15874 is the only specimen having 
the third vertebral longer than wide. 

The most distinctive feature of the plastron is the extended epiphis- 
tnd lip that always projects well Ix'vond the line of the front margin of 
the carapace. It was the "marked prominence and size of the 
epiplastral lip" that Lanibe' stressed as the most important character 
for distinguishing Testvdo praeextans, a character to which the specific 
name refers. In the light of these additional spc'cimens it is clearly 
shown that the extent and shape; of this lip constitute one of the more 
stable characters of this species. 

< I-ambp. L. M., Ottawa Nat., vol. 27, p. W, 1013. 

• LanilH!, L. M., Ottawa Nat., vol. 27, pp. 57-fil, pl«<, 1. o. UM.1. 



302 



PROCEEDINGS OF THE NATIONAL MUSEUM 



The relative proportions of the lobes and other measurements of the 
plastron are clearly shown in table 6. 

Table 4. — Comparative measurements (in millimeters) of neurals of Testudo 

praeextans 



Specimen 


Length 


Width 


1 


2 


3 


4 


5 


6 


7 


8 


9 


1 


2 


3 


4 


5 


6 


7 


8 


9 


C. N. M. No. 8401 (type) 

U.S.N.M. No. 15878 

U.S.N.M. No. 16732 

U.S.N.M. No. 15874 

U.S.N.M. No. 16728 


68 
62 
67 
76 
76 


40 
42 
46 
48 
43 


41 
37 
43 
44 
42 


37 
39 
41 
39 

54 


36 
36 
55 
42 


34 
34 
36 
34 


41 
38 
35 


15 
46 
51 


38 


44 
33 
34 
46 

48 


47 
42 
49 
51 
55 


34 
40 
41 
39 
43 


48 

49 
42 
54 


45 

45 
51 


44 
50 

58 
46 


44 
49 
57 
55 


35 
38 
53 


39 



Table 5. — Comparative measurements (in millimeters) of vertebrals of Testudo 

praeextans 



Specimen 


Length 


Width 


1 


2 


3 


4 


5 


1 


2 


3 


4 


5 


C. N. M. No. 8401 (type) 

U.S.N.M. No. 15878 

U.S.N.M. No. 16728 _ _ 


100 
100 
103 
120 
122 


77 
82 
91 
97 
109 


78 
81 
86 
90 
112 






97 
110 
113 
114 
100 


82 
79 
91 
91 
86 


88 

86 

103 

101 

90 


82 

87 


16? 


103 


105 


170 


U.S.N.M. No. 16732 

U.S.N.M. No. 15874 


109 

86 


106 
117 


97 
89 


211 

188 



Table 6. — Comparative measurements (in millimeters) of plastra of Testudo 

praeextans 



Measurement 


C.N. M. 
No. 8401 


U.S.N.M. 
No. 15878 


U.S.N.M. 
No. 16732 


U.S.N.M. 
No. 15874 


U.S.N.M. 
No. 16728 


Greatest length of plastron 

Length of anterior lobe 

Width of anterior lobe . 


493 
163 
229 
147 
239 
72 
97 


495 
154 
205 
135 
225 
60 
81 

47 
34 


540 
178 
258 
145 
245 
71 
95 

72 
39 


580 
187 
243 
155 
270 
85 
102 

88 
48 


191 


Length of posterior lobe 

Width of posterior lobe 

Length of epiplastrai lip 

Width of lip at base , _ 


76 
97 


Extension of epiplastrai lip be- 
yond border of carapace. _ 


75 


Depth of posterior notch 


40 





The free borders of the plastra in all five specimens are acute, 
except those parts of the borders adjacent to the notches, which 
are thickened and rounded. The posterior lobes are terminated 
behind in two broadly rounded apices separated on the median line 
by wide V-shaped notches. The borders of the apices are usually 
slightly toothed. Likewise, the anterior ends of the epiplastrai lips 
are notched at the center and the borders are toothed with six blunt 
teeth, except in the type of Testudo praeextans, as shown in figure 25. 



OSTEOLOGY OF TESTIDO PHAEEXTANS — GILMORE 303 

The entoplastra in all the specmiens except No. 15878 are in 
agreement in having pointed anterior ends and a broad somewhat 
rounded posterior border, with the gular scutes overlapping the 
front of this bone. 

There is some variation in the extent of the plastral scutes in these 
specimens, as shown in table 7. 

Pelvis. — The pelvis of specimen No. 15874 is in an unusually perfect 
state of preservation, the two halves being coalesced along the median 



Figure 25. — Plastron of Testudo praeexlans Lambc, type (C. N. M. No. 8401): ab, abdominal 
scute; an, anal scute; ent, entoplasiron; /^m, femoral scute; g, gular scute; hum, humeral 
scute; hyo, hyoplastral bone; hypo, hypoplasiral bone; pec, pectoral scute; xiph, xiphi- 
piastral bone. One-sixth natural size. 

line, as reprcscntt-d in figures 20 and 27. The i)elvis of No. 107i{2 
is also well preserved, but the two halves were separated along the 
median suture. Except for minoi- details these two pelves ai-e in 
close agreement and thus are probably tyi)ical of Testudo pratcxtaiis. 

Of these two turtle specimens of nearly equal size and presumably 
of equivalent age om(> has the pelvic sutures coalesc(>d, while in 
the other these same sutures are open, thus indicating that the 
relative age of an individual cannot always be judged by the non- 
coalescence of the sulural junetures. 

The anterior extremities of the pui)es are complete in specimen 
No. 15874, which shows that border to be broadly rounded, thus 



304 



PROCEEDINGS OF THE NATIONAL MUSEUM 



suggesting that Hay may have erred m restoring this missmg end 
in the pelvis of the type of T. laticunea as being bilobed.® The un- 




FiGURE 26. — Pelvis of Testudo praeextans Lambe (U.S.N.M. No. 15874), viewed from above: 
il, ilium; is, ischium; pu, pubis. One-half natural size. 




Figure 27. — Pelvis of Testudo praeextans Lambe (U.S.N.M. No. 15874), viewed from the 
left side: il, ilium; is, ischium; pu, pubis. One-half natural size. 

usual length of the anterior pubic process, forward of the lateral 
pubic tuberosities, appears to be a distinctive feature of this species. 
The apices of the ischial tuberosities are bluntly pointed and 80 mm. 
apart. 

« Hay, O. P., Carnegie Inst. Washington Publ. 75, p. 404, flg. 512, 1908. 



OSTEOLOGY OF TESTUDO PRAEEXTANS — GILMORE 



305 



Careful comparison made of the other skeletal parts preserved 
with these various specimens, with each other and with homologous 
bones of other species of Testudo, failed to disclose, except that of 
size, differences that would be of assistance in specific differentiation. 
On that account there seems do point in illustrating or describing 
these parts here, although most of the bones present are in excelhMit 
preservation. In order, however, that there may be a record of rela- 
tive proportions between carapace and limb bones, a table of measure- 
ments of the more important elements of the specimens here dis- 
cussed is given (table 8). 



Table 7. 



-Comparative measurements {in millimeters) of plastral scutes of Testudo 
praeextans 



Scute flength on the midline) 


C.N.M. 

No. 8401 

(type) 


U.S.N.M. 
No. 15878 


U.S.N.M. 
No. 16732 


U.S.N.M. 
No. 15874 


U.S.N.M. 
No. 16728 


Gular - 




93 


80 
103 

43 
147 

58 

46 


162" 
44 

143 
72 
51 


92 


H umeral 






Pectoral 

Abdominal 


34 


135" 
63 
50 




Femoral 






Anal 













Table 8. — Comparative measurements (in millimeters) of girdle and limb bones of 

Testudo praeextans 



Measurement 



U.S.N.M. 
No. 15878 



U.S.N.M. 
No. 16732 



U.S.N.M. 
No. 16728 



U.S.N.M. 
No. 15874 



Greatest lenp;th of coracoid 

Greatest width at inner end 

Greatest length of scapula from tip to tip. 

Greatest length of humeru.s 

Greate.'^t length of ulna 

Greatest length of radius 

Greatest length of femur 

Greatest length of tibia. 

Greatest lengt h of fibula 



80 



88 
77 
153 
155 
99 
98 



82 

65 

151 

153 



78 



77 
80 



74 

76 

1 150 

157 
99 
97 

113 
82 
82 



I Estimated. 



Feet. — Ail four of the National Museum specimens considered in the 
I)receding pages have various parts of the carpi, tarsi, and feet present, 
l>Mt only in No. 15878 arc the bones preserved at all in articulation. 
Tlie right forefoot, with cari)us and radius, although lacking a few 
elements, has the others articulated in normal relationship as siiown in 
plate 44, figure 2. This foot clearly shows the presence of five well- 
developed digits, each having a daw. The snudl blocklike bones of 
the carpus are but little disturbed and can be quite certainly identified. 
The ulnare and intermedium furnish the main articulation of the ulna, 
which is missing. Occujjying the position of the radiale at th(> distal 
end of the radius is an angular blocklike element that, following Baur 



306 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

and Williston/ we may identify as centrale 1. Its outer side is in 
juxtaposition to a smaller pentagonal bone regarded as centrale 2. 
These two centralia although distinct elements in this individual are 
often found fused into a single bone, and this appears to have happened 
in specimen No. 15874. Williston, in the book cited, observes/' Among 
terrestrial tortoises the radiale has disappeared until nothing is left of 
it but a nodule of cartilage united with the first centrale which has 
usurped its place." Hay, however, in his "Fossil Turtles of North 
America," continues to regard the first centrale as the radiale. 

The fifth carpale (pi. 44, fig. 2, C5) is present and remains in 
articulation with metacarpal 5. Carpalia 3 and 4 are missing. The 
third digit lacks its metacarpal, and a phalange is missing from the 
fifth digit; otherwise the foot is complete. Comparison of these 
wrist and foot bones with those of the living Galapagos Testudo of 
comparable size shows a close correspondence in form, as well as in the 
arrangement of the individual elements. 

The hind foot of No. 15878 contributes nothing new to our knowl- 
edge of the pes of Testudo. 

SUMMARY 

In the original description of Testudo praeextans Lambe called 
attention to the close similarity in the form of the epiplastral beak to 
that of T. thomsoni but concluded that the "much greater propor- 
tionate size of the epiplastral lip and differences throughout of the 
elements forming the lobe" were sufficient to indicate their specific 
distinctness. 

In view of the very fragmentary character of the type of T. thomsoni, 
and especially of the considerable variation in the form of the anterior 
lobe as shown by the present specimens, it would now appear that 
Lambe was not justified in establishing the new species T. praeextans. 
On plastral parts alone I should unhesitatingly regard T. praeextans to 
be a synonym of T. thomsoni, which has priority by several years. 
However, when the skulls are compared, differences in proportion of 
the elements forming the skull roofs, and the different widths of the 
channels on the triturating surfaces, strongly suggest that the dis- 
covery of more complete materials of T. thomsoni may disclose other 
and more important distinctive characters. For the present, there- 
fore, it seems desirable to continue the use of both names. 

In the preliminary study of the present materials it was first 
thought that these specimens could not be specifically distinguished 
from Testudo laticunea Cope and that such differences as existed might 
be attributed to the female sex of the type of that species. This idea 
was abandoned, however, with the discovery that the Wyoming 

' WUliston. S. W., The osteology of the reptiles, p. 179, 1925. 



OSTEOLOGY OF TESTUDO PRAEEXTANS — GILMORE 307 

spociinons all hail i)it)aclly roundtil aiitoiior carapace borders and 
prominent epiplastral lips with parallel sides, as contrasted with the 
more truncate carapace hoidi^r and wedge-shaped epiplastral lip in 
T. lat'ivuiua. 

This study of a number of Testudo specimens from a restricted area 
antl apparently of a single species shows enough variation in shell 
structure to cast much doubt on the validity of many described species, 
especially those foundetl on fragmentary materials. Furthermore, it 
raises the perplexing question as to what featvn'es are to be relied on 
for specific difrcrentation. It has been demonstrated (pis. 38-41) 
that the neural, costal, and other elements forming the carapace, as 
well as the form and proportions of the vertebral and plastral scutes, 
are seldom in themselves suflicientl}' constant in shape and proportion 
to be relied on for specific designation. 

^^^lether characters of the skull in fossil Testudo will be found more 
stable can only be determined when a series of crania is available for 
comparison, and at this time no such series exists. It is in the general 
form of the carapace and plastron and especially in the development 
of epiplastral beak that most reliance has been placed in diagnosing 
the present species. 

TESTUDO LATICUNEA Cope 

Plates 42, 43 

Testudo lalicunea Cope, Paleont. Bull. No. 15, p. 6, 1873. — Hay, Carnegie Inst. 
Wa.shington Publ. 75, p. 402, pi. 67, figs. 1, 2, 1908. 

Two specimens (U.S.N. M. Nos. 15854 and 16731) in the Oligocene 
collections from eastern Wyoming are identified as pertaining to 
Testudo laticunea Cope, the first recorded occurrence of the species 
in this area. These specimens are remarkably alike (cf. pis. 42 and 
43) both in size and structural detail, a condition calling for comment, 
as anyone will agree who has had occasion to study a series of Testudo 
specimens. 

The type of T. laticunea is an essentially complete shell, with some 
of the internal skeleton, and was found in the Oligocene badlands 
along the head of Horse Tail Creek in northeastern Colorado. Two 
other specimens, one from the Tiianotherium beds (Chadron) of South 
Dakota and one from the Oreodon beds (Brule) of Siou.x County, 
Nebr., were identified by Hay * as jx'rtaining to this species. These 
together with the two in the U. S. National Museum comi)rise all 
the known materials of the .species. 

Specimen U.S.N.M. No. 15854 consists of a nearly complete cara- 
pace and plastron (pi. 43), left humerus, proximal end of right ; left 
ulna, radius, and part of forefoot; incom])letc right scai)ula, left 

' Hiiy, o. v., CamoKie Inst. Wa.shlDKlon I'ubl. T.'i, \t\). IOt-40:., 1908. 



308 



PROCEEDINGS OF THE NATIONAL MUSEUM 



coracoid; left femur, tibia, fibula, and portion of hind foot. It was 
collected by C. W. Gilmore on the Thomas Ranch, Niobrara County, 
Wyo., in 1932. Specimen U.S.N.M. No. 16731 consists of a complete 
carapace and plastron (pi. 42). It was collected 1 mile northeast of 
Whitman Postoffice, Niobrara County, Wyo., by George B. Pearce 
in 1942. Both of these specimens are from the Brule division of the 
Oligocene. 

In size and general contour of the shell these specimens are in close 
agreement with the type, as shown by the measurements given in 
table 9. 



Table 9. 



-Comparative measurements {in millimeters) of carapace and plastron of 
Testudo laticunea 



Measurement 



Type spec- 
imen 



U.S.N.M. 
No. 15854 



U.S.N.M. 
No. 16731 



Greatest length of carapace _■ 

Greatest width of carapace 

Greatest length of plastron 

Gi-eatest length of anterior lobe 

Greatest width of anterior lobe 

Greatest length of posterior lobe 

Greatest width of posterior lobe 

Greatest width of bridge 

Greatest width of anterior lip at gUlar notch. 



408 
356 
440 
132 
200 
120 
200 



87 



436 
332 
440 
132 
202 
110 
193 
200 
90 



435 
340 
443 
137 
203 
110 
196 
197 



Specimens U.S.N.M. Nos. 15854 and 16731 are practically free from 
distortion and thus give a true picture of the normal shell. It will be 
noted in the table of measurements that the Wyoming carapaces are 
considerably narrower than that of the type. This difference may be 
partly due to the crushing to which the type has been subjected. In 
cross section at midlength the shells of Nos. 15854 and 16731 are evenly 
rounded from side to side, whereas the type is said to be flattened on 
top. All three, however, can be classed as having a depressed style of 
shell, and all are of female sex, as shown by the flatness of their plastra. 

Other minor differences observed between these two specimens and 
the type fall well within the variations expected in individuals of a 
single species. 

The skeletal parts preserved with the shell of U.S.N.M. No. 15854 
have been carefully compared with the homologous bones of T. 
praeextans and other species of Testudo, but except for their smaller 
size no other characters for distinguishing between them were dis- 
covered. 

Hay ^ has pointed out in his study of the type of T. laticunea that the 
"neurals have not attained so liigh a degree of differentiation as they 
have in most species of the genus." Although the complete neural 
series cannot be traced out in either specimen, the correctness of the 



» Hay, O. P., Carnegie Inst. Washington Publ. 75, p. 403, 1908. 



0STE01.0(.Y OF TESTUDO PUAEEXTANS — GILMORE 



309 



above coju'liision is verified ])y these new materials. S])eeinien No. 
16731 np])enrs to sliow the seeoiitl neural to he oetagoual as it is in the 
South Dakota specimen studied hy Hay.'' 

The slight variation in the vertehrals of the three s])eeiniens dis- 
cussed here is shown by the measurements given '\u table 10. 

Table 10. — Comparative measurements {in millimeters) of veriebrals o/ Testudo 

laticunea 



Specimen 




Length 






Width 




1 


2 


3 


4 


5 


1 


2 


3 


4 


s 


Type specimen 


86 
78 
91 


80 
89 

77 


81 
83 
78 


75 
83 
78 


90 
88 
90 


102 
94 
94 


80 

82 
74 


85 
82 
78 


82 
'79' 


1 :',:, 


U.S.X.M. No. 15854 

U.S.N. M. No. 16731 


133 
142 



Table 11. — Comparative measurements (in millimeters) of epiplastral beak of Tes- 
tudo quadra ta 



Measurement 


Type 
(A.M.N.H.) 


U.S.N. M. 
No. 16737 


Width of beak at base . . 


120 
70 
29 


135 


Length of beak from gular groove 


54 


Thickness of beak at base . 


41 







The combination of characters that appear to distinguish Testudo 
laticunea Cope is as follows: 

Diagnosis: Carapace depressed with truncated anterior border, but 
slightly rounded from side to side across the vertebral region; prom- 
inent epiplastral lip, projecting beyond the borders of the carapace 
with converging sides; neurals less diU'erentiated than in most species 
of the genus, and usually w^ithout those of octagoiuil form. 

TESTUDO QUADRATA Cop* 
PlATK 44, FiGURK 1 

Testudo quadratus Cope, The V'ertebrata of the Tertiary formations of the West, 
p. 704, pi. 01, fiK. 5, 1884. 

Testudo quadrata Hay, HiblioKraj)hy ami catalogue of the fo.s.^il V'ertebrata of 
North America, p. 451, 1902; CarneRJe Inst. Washington I'ubl. 75, p. 410, 
figs. 532, 533, 1908; Seconrl bibliography and catalogue of the fossil Verte- 
bra ta of North America, vol. 2, p. 104, 1930. 

A third species of 7'€.s<7/(/o occurring in the Indian Creek Hiisin area 
is represented in the National Museum OligoccMu- ((tllcction by an 
e])iplastral beak (U.S.N.M. No. 10737), shown in ])\iiiv. 44, figure 1. 
This specimen was collertcd from the Brule, aixiut 1 mihr north of 
W hitman Postoflice, Niobrara County, Wyo., in 1942. 

» Ibid., p. 4fH. 



310 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

The quadrate form of this beak with diverging lateral edges is more 
like that of T. quadrata Cope than of any other species, but it differs 
in having the gulohunieral sidci running backward and inward to 
meet on the median line instead of directly across this bone at right 
angles to the midline, as in the type of the species. This feature, 
stressed by both Cope and Hay, is so at variance with all other known 
species of Testudo, both living and extinct, that it leaves one wondering 
if it is not an abnormal condition peculiar only to that individual. 

Specimen No. 16737 is larger than the type, as shown by the 
measurements given in table 11. 

With the possible exception of length, the other differences observed 
between the two specimens may be regarded as individual variations. 
The beak is shorter than that of the type although exceeding it in all 
other dimensions. Regardless of the question of doubt that this 
difference may engender as to its proper specific assignment, this 
fragmentary specimen certainly indicates the presence in the Brule 
of the Hat Creek Basin of a third species of Testudo. 

NOTE ON THE GEOLOGICAL DISTRIBUTION OF TESTUDO 
IN NORTH AMERICA 

In the "Fossil Turtles of North America," p. 397, 1908, Hay lists six 
species of Oligocene Testudo as occurring in the Oreodon beds (Brule) 
and two as coming from the Titanotherium beds (Chadron), as follows: 

Brule Chadron 

Testudo amphithorax Cope T. brontops Marsh 

T. ciiltrnta Cope T. exornata Lambe 

T. laticunea Cope 
T. ligonia Cope 
T. quadrata Cope 
T. ihomsoni Hay 

In a subsequent publication, "Second Bibliography and Catalogue 
of the Fossil Vertebrates of North America," Hay assigned all the 
Brule species to the Chadron. One is at a loss to understand whether 
this change was made on the basis of new information or whether it was 
a slip of the pen. 

In view of the recognition of two, if not three, of the species in im- 
doubted Brule deposits, it would appear to show the original geologic 
assignment was correct. That some of these species may have per- 
sisted from the Chadron into the Brule is suggested by a specimen in 
the Yale Aluseum from the Titanotherium beds, which Hay '^ identified 
as belonging to Testudo laticunea. That such was the case needs ad- 
ditional verification. Tliis assertion is made on the fact that the 
commonest of all land tortoises of the Brule, Stylemys nebrascensis, has 
never been recognized as occurring in the Chadron. 

11 Hay, O. P., Carnegie Inst. Washington Publ. 75, p. 404, 1908. 

U. S. GOVERNMENT PRINTING OFFICE: 1946 



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SMITHSONIAN INSTITUTION 

U. S. NATIONAL MUSEUM 
VoL % Wuhington: 1946 No. 3200 

EIGHT NEW SPECIES OF CHALCID-FLIES OF THE GE^^US 
PSEUDAPHYCUS CLAUSEN, WITH A KEY TO THE 
SPECIES 



By A. B. Gahan 



The species of chalcid-flies of tlie genus Psevdaphycus Clausen 
(family Encyrtidae) are apparently all parasites of pseudococcine 
scale insects, and some of the species are known to be of considerable 
economic importance through the control they exercise over their 
hosts. P. utilis Timberlake is credited with having all but exter- 
minated Pseudococciis nipae (Maskell) in Hawaii, and the new species 
malinius (described on a subsequent page of this paper), which was 
introduced into the eastern United States from Japan to combat 
Pncudoroccu.<i cowHtocki (Kuwana), apparently gives promise of 
bringing that serious orchard pest under control. 

Tlie genus is widely distributed. Species are now known to occur in 
Austria, Spain, Canary Islands, Cuba, Puerto Kico, the United States 
from New Jersey to California, Hawaii, Korea (Cho.sen), and Japan. 

PypAidaphyffUA belongs to the group of genera related to Aphycus 
Mayr. It may be distinguished from Aphycus as well as most of the 
other related genera by the fact that it has only live, instead of six, 
segments in the antennal funicle. It is most closely related io Both- 
ri^orcwra Timberlake and A<-e,ropha(ni.f< Smith, both of which have 
the funicle 5-s('grnented. It may be separated from linthriocrarra 
by the dull sculpture of its frontovertex and tlioraric dorsum and 
by the fact that it is never shining black in color. As pointed out by 
Mercet (liull. Knt. Kes., vol. 2H, p. ;H7, 1!»;57), the dillerences between 
Pftetidaphycus and AcerophnguJi are very slight. About the only real 
differences seem to l)e that in Arerophng'>t-f( the front(»vertox is less than 

311 



705901— 4e 



312 PROCEEDINGS OF THE NATIONAL MUSEUM tol. 9e 

twice as long as wide, the ocellar triangle is more or less obtuse, and 
the antennae are unicolorous and always pale, while in Pseiidaphycus 
the f rontovertex is nearly always two or more times as long as broad, 
the ocelli are arranged in an equilateral or acute triangle, and the 
antennae are always contrastingly colored with the scape, pedicel, 
and at least some of the funicular segments black or fuscous and the 
club white. 

The species of Pseuda'phycus offer very few good characters for 
their separation. Slight differences in sculpture are hard to define 
and differences in color consist largely of differences in shades of yel- 
low, since all the species are yellowish sometimes varied with admix- 
tures of blackish or fuscous. The following key to species is offered 
as an aid to identification but should be supplemented by comparison 
with types or authentically determined specimens whenever possible : 

KEY TO THE DESCRIBED SPECIES OF PSEUD APHYCUS 

FEMALES 

1. Pedicel of antenna as long as or longer than first three segments of funicle 

combined, funicular segments from first to last successively increasing in 

width and all more or less transverse 2 

Pedicel of antenna not longer than first two funicular segments combined; 
first funicular segment small, transverse, narrower than pedicel ; segments 
2 to 5 of funicle subquadrate, subequal, and each a little broader than 
pedicel 1. meritorius, new species 

2. Ovipositor exserted one-fifth to nearly one-half length of abdomen 3 

OvijDOSitor extending barely beyond apex of abdomen, at most distinctly less 

than one-sixth length of abdomen 2. prosopidis Timberlake 

3. Wings hyaline; scape not expanded beneath 4 

Wings at least faintly fuscous ; scape often but not always somewhat thick- 
ened or slightly expanded beneath 5 

4. General color pale lemon-yellow ; hairs on mesoscutum white; ocellar triangle 

rather small and nearly equilateral; ovipositor exserted about one-third 

length of abdomen 3. graminicola Timberlake 

€reneral color bright orange-yellow; hairs on mesoscutum dark brown or 
blackish ; ocellar triangle obviously acute and unusually large ; ovipositor 
exserted about one-fifth length of abdomen 4. utilis Timberlake 

5. Porewing fuscous with a more or less distinct paler hyaline or subhyaline 

transverse streak beyond apex of stigmal vein; scape obviously a little 

'''' thickened or broadened 6 

S'',Forewing uniformly fuscous or subfuscous, without a paler transverse streak; 

,.v scape usually slightly thickened but sometimes cylindrical 9 

<5. Anterior ocellus at center of f rontovertex ; speculum of forewing not inter- 
rupted; mesoscutum, axillae, and scutellum bright orange-yellow, with a 
conspicuous narrow transverse band of black at anterior margin of meso- 
scutum 5. websteri Timberlake 

Anterior ocellus distinctly above center of f rontovertex ; speculum of forewing 
interrupted a little behind middle ; mesoscutum, axillae, and scutellum more 
or less dusky orange to blackish yellow, anterior margin of mesoscutum with 
suture sometimes blackish but never forming a conspicuous and well- 
defined transverse band 7 



CHALCID-FLIES OF GENUS PSEUDAPHYCUS — GAHAN 313 

7. Mesoscutum, axillae, scutellum, and abdomen blackish yellow; lateral borders 

of mesoscutum white; posterior ocelli as far from anterior ocellus as from 
margin of occiput; Inner orbits parallel for whole length of froritovertex. 

6. maculipennis Mercet 

Mesoscutum, axillae, and scutellum dusky orange-yellow; lateral borders of 

mesoscutum not white; posterior ocelli a little more distant from anteriof 

ocellus than from margin of occiput; inner orbits diverging very slightly 

below 8 

8. Frontovertex about two and one-half times as long as broad ; head and thorax 

dorsally rather pale orange-yellow, axillary sutures of mctanotum usually 

not blackish 7. angelicus (Howard.) 

Frontovertex fully three times as long as broad; head and thorax dorsally 
a deeper shade of orange-yellow mixed with blackish, and axillary sutures 
brownish black 8. abstrusus, new species 

9. Middle and hind tibiae each with two fuscous or blackish bands 10 

Middle and hind tibiae without fuscous bauds 12 

10. Frontovertex as broad as long or somewhat broader than long; ocelli in an 

equilateral triangle; dorsum of thorax dirty yellow; antenna fuscous with 

first funicular segment and club white 9. austriacus Mercet 

Frontovertex twice to two and one-half times as long as broad ; ocellar triangle 
at least slightly acute; dorsum of thorax orange-yellow, antennal scape 
yellowish below, brown above; pedicel and funicle brownish or fuscous, 
fifth funicular segment and club white 11 

11. Ocellar triangle distinctly acute, posterior ocelli about half as far from each 

other as from anterior ocellus ; speculum of forewing interrupted below 

middle 10. orientalis Ferrlere 

Ocellar triangle nearly equilateral ; posterior ocelli only slightly closer to 
each other than to anterior ocellus ; speculum of forewing not Interrupted. 

11. malinus, new species 
12 Last two segments of funicle (sometimes entire funicle) and club white, 
first three segments of funicle usually fuscous; frontovertex unusually 
narrow, fully three times as long as broad ; ocelli in a very acute triangle, 
posterior pair separated by about diameter of an ocellus ; ovipositor one- 
third to half as long as abdomen 12. angustifrons, new species 

Entire funicle black or at least fuscous; frontovertex at least a little broader; 
ocellar triangle less acute and posterior pair of ocelli separated by more 
tlian diameter of an ocellus; ovipositor usually shorter 13 

13. Mf?«o8cutum, scutellum, and axillae weakly sculptured, distinctly shining; 

dorsum of thorax dirty yellowish or grayish in color ; antennal club blackish 

basal ly 14 

Mesowutura, scutellum, and axillae (Wiely and densely Hcul[)tured, dull ; dorsum 
of thorax orange-yellow; antennal club entirely white 15 

14. Scape of antenna obviously a little thickened or expanded beneath ; ovipositor 

exserted one-third h'ngth of abdomen ; 8p<'culuni of fonnviug uninterrupted. 

13. mundus, new sp«'cle»* 

Scape of antenna not obviously expandt-d ; f)vlposltor about on<!-!lfth length 

of abdomen ; speculum at least partially Interrupted a little behind middle. 

14. liinatulus, n«'w six-cles 

15. Anterior margin of face narrowly dark brown or blncklHh, forming a dark 

transverse line between bases of mandibles; occiput exct'pt around margins 
black or blackish, anterior face of pronotum mostly black; suture between 
pronotum and meso.scutum blackish; tegula pale at base, blackish aplcnily; 
abdomen dorsally mostly blackish; posterior tibiae often with two fusrons 
bands 11. malinus, new siM'cles 



314 PROCEEDINGS OF THE NATIONAL MUSEUM vol. »« 

Anterior margin of face without a dark line between bases of mandibles; 
head and thorax dorsally bright orange-yellow without any blackish mark- 
ings; tegula white; abdomen with four rather definite dark transverse 
bands, one of which connects vibrissal plates, one basad of this and two 

apicad of it; hind tibiae not banded 16 

16. Frontovertex bright orange-yellow and very finely granulosely sculptured 
with only very faint indication of alveolation even under high magnifi- 
cation 15. meracus, new species 

Frontovertex paler, dilute orange-yellow, entire surface covered with fine 
but very distinct facetlike punctures 16. alveolatif rons, new species 

1. PSEUDAPHYCUS MERITORIUS, new species 

The subquadrate second to fifth segments of the f unicle distinguish 
this species from all other species treated here. 

Female. — ^Length 0.92 mm. Frontovertex, mesoscutum, axillae, 
and scutellum slightly dusky orange-yellow ; head (except frontover- 
tex), tegulae, pronotum, all pleura, and sterna and all legs yellowish 
white; propodeum fuscous; abdomen dorsally yellowish with four 
transverse fuscous bands, the anterior two bands coalesced at the 
lateral margins of abdomen and enclosing an ellipsoidal area between 
the cerci, the posterior two bands straight ; abdomen beneath fuscous 
to blackish medially with the margins pale, ovipositor pale at base, 
darker apically ; eyes black ; ocelli dark red ; antennal scape pallido- 
fuscous; pedicel and entire f unicle black or blackish; club white; 
wings uniformly faintly fuscous, or subhyaline ; marginal and stigraal 
veins dark fuscous, submarginal vein pale. 

Frontovertex a little less than twice as long as broad, granulosely 
punctate; inner orbits parallel; ocelli in an equilateral triangle; 
anterior ocellus situated only slightly above middle of frontovertex; 
lateral ocellus less than its own diameter from margin of eye and 
about its own diameter from occipital margin ; eyes clothed with short 
hairs. Antenna clavate; scape not expanded beneath, five to six 
times as long as thick; pedicel nearly twice as long as thick and less 
than one-third as long as scape; first funicular segment small, nar- 
rower than pedicel ; segments 2 to 5 of f unicle subequal and subquad- 
rate, and each a little thicker than pedicel; club a little thicker than 
funicle and aipproximately as long as the four preceding funicular 
segments combined. 

Mesoscutum, axillae, and scutellum finely granulosely punctate, 
mat; propodeum in front of spiracle weakly reticulated, elsewhere 
smooth ; pleura weakly reticulated, slightly shining. Forewing with 
nearly uniform, rather dense discal ciliation ; speculum closed at pos- 
terior margin of wing but otherwise uninterrupted; marginal vein 
punctif orm ; postmarginal vein shorter than stigmal. 

Abdomen short-ovate in outline, about as long as thorax or slightly 
longer, slightly narrower than thorax ; weakly reticulately sculptured 



CHALCID-FLIES OF GENUS PSEUDAPHYCUS GAHAN 315 

and more or less distinctly shining dorsally and with similar sculpture 
vent rally. Ovipositor exserted one-fourth to one-third the length 
of abdomen. 

}fale. — Length 0.8 mm. Similar to female except as follows: 
Funicular segments subequal in length, but successively increasing in 
width from first to last, the fifth about as broad as pedicel and a little 
broader than long; club nearly as long as funicle; abdomen shorter 
than thorax, nearly circular in outline, blackish dorsally and ventrally 
with yellowish lateral margins above and beneath except at cerci, 
where the black on dorsum extends to the margins. 

Type locality. — Winchester, Va. 

Type.—V. S. X. M. No. 57324. 

lie marks. — Described from numerous specimens received from 
D. W. Clancy of the Charlottesville, Va., laboratory of the Bureau of 
Entomology and Plant Quarantine and said to have been reared from 
Fseudococcus virgatus (Cockerell) collected at Winchester, Va., June 
20, 1944, and recorded under laboratory note Nos. 67G, 677, 678, and 
681. Other specimens identified as this species were reared from the 
same host collected at Winchester, May 31, 1944, under laboratory No. 
638; August 4, 1943, laboratory No. 529; August 3, 1944, laboratory 
No. 850; and at Hamilton, Va., August 12, 1943, laboratory No. 530, 
and May 30, 1944, laboratory Nos. 634 and 635. 

2. PSEUDAPHYCUS PROSOPIDIS Timberlake 
P-< udaphyrus prosopidin TiMiJFmL.\KK, Proc. U. S. Nat. Mus., vol. 50, p. 571, 1916. 

Described from Mesilla, N. Mex., as a parasite of Fseudococcus 
prosopidh ( Cockerell ) . 

I. PSEUDAPHYCUS GRAMINICOLA Timberlake 

Pfifudaphyrus graminUola Ti.mbeklake, Proc. U. S. Nut. Mus., vol. 50, p. 570, 1916. 

Kecorded by Timberlake as parasitizing pseudococcine scales on 
tStipa sp. at Las Vegas, N. Mex., and on Elymus condctu'<atus at Kim- 
balls, Utah. The host insect involved in the record from Stipa sp. 
has recently been ideiifified by Aforrison as Trionyiuu.s ncomcxicamus 
var. utahercHis (Cockerell) . The host on fJhjnius remains unidentified. 

4. PSEUDAPHYCUS UTILIS TlmbcrUke 

pMMila^hycuH ufiliH TiMfJE:Ri..\KK, proc. Hiiwriiiaii Kiit. Soc, vol. ."">, p. .'{23, Akh. 1-4, 
lfrj3.— SwcETi', Pr.tc. Hiiwaiiiui Kiit. Soc. vol. .1, p. »U, 11)2:{. — Ti.MitFniLAKK, 
Proc. llawaiiiiri Eiit. Soc., vol. .""), p. 431. WS.i. — Williams. IlMinlborik of iii.vtrts 
Jiiid other iiiv(Ttcl>ratt's of Hawaiian su^j.-ir cane flchlH. ]). L'ri.'i, lll.'n.- Swkzky, 
Pror. nth Pafiflc Sol. Con^rr. (Canada, 19:{.3), vol. .'>, [». rj.WJ, H>34.— PrMBianoN, 
Hawaiian Planters Hoc, vol. 45, p. lOO, 1J>41. — Anon., Puerto Hico Kxp. Stat. 
Hrj.t. for imo. p. 68. lt)42. 

This species is a parasite of PseudococoiM nipae (Maskell) and was 
originally described from the state of Veracniz, Mexico. It was intro- 



316 PROCEEDINGS OF THE NATIONAL MUSEUM vol. w 

duced into Hawaii in 1922 where, according to Pemberton, it proved so 
efficient as practically to exterminate its host. More recently it was 
introduced into Puerto Rico where, according to the above-cited report 
of the Puerto Eican Experiment Station, it gives every indication of 
bringing the host insect under complete control. 

5. PSEUDAPHYCUS WEBSTERI Tiraberlake 
Pseudaphycus wetsteri Timberlake, Proc. U. S. Nat. Mus., vol. 5, p. 570, 1916. 

Known only from the type, a single female, said to have been reared 
from the stems of Elyrrms virginicus, at Villa Ridge, 111. The real 
host was probably a pseudococcine scale on the Elymus, as Timber- 
lake pointed out. 

6. PSEUDAPHYCUS MACULIPENNIS Mercet 

Pseudaphycus macuUpennis Mebcbt, Bol. Real Soc. Espan. Hist. Nat, voL 23, 
p. 140, fig. 2, 1923; Rev. Espan. Ent., vol. 1, p. 12, fig. 1, 1925. 

Originally described from material collected on the island of Tene- 
riffe, Canary Islands. Mercet later collected it at Barcelona, Spain. 
No host has as yet been recorded for the species. 

7. PSEUDAPHYCUS ANGELICUS (Howard) 

Aphycus angelicus Howard, Proc. U. S. Nat. Mus., vol. 21, pp. 241, 245, 1898. 

Pseudaphycus angelicus (Howard) Timberlake, Proc. U. S. Nat. Mus., vol. 50, 
p. 573, 1916.— Clausen, Univ. California Techn. Bull., Entom., vol. 3, pp. 258, 
280, 1916.— Flanders, Journ. Econ. Ent., vol. 28, p. 552, 1935 ; vol. 33, p. 758, 
1940. 

P. angelicns was originally described from Los Angeles, Calif., as 
a parasite of Pseudococcus sp. on passionflower. It has since been 
recorded by Timberlake as parasitizing Pseiuiococcus {longispinus 
Tsirgiom) = adonidum (Linnaeus), ryani (Coquillett), and {citro- 
philus Clausen) = gahani Green; by Clausen from Pseudococcits 
maritiinus (Ehrhorn) and gahani Green; and by Flanders from 
Pseudococcus adonidu/m (Linnaeus) and Phenacoccus gossypii Town- 
send and Cockerell. All these records are of occurrences of the species 
in California, and thus far it is not known to occur outside of that 
State. 

Timberlake has given a satisfactory description of this species in 
the paper cited above, except that it should be not^d that the trans- 
verse fuscous band on the cheeks which he mentions is not always 
present. 

8. PSEUDAPHYCUS ABSTRUSUS, new species 

The specimens to which this name is here assigned have proved 
exceedingly puzzling (hence the specific name). 

Female. — Length 0.94 mm. Apparently agreeing in every respect 
with females of angelicus except that the frontovertex is very slightly 
narrower than in that species, and the general color is somewhat 



CHALCID-FLIES OF GENUS PSEUDAPHYCUS GAHAN 317 

tiarker, dirty orange-yellow, with the anterior margin of mesoscutum 
as well as the axillary sutures blackish and the metanotum, propodeum, 
and entire dorsum of abdomen deeper black. The color of antennae, 
uudeipnrts of the head and thorax, tegulae, wings, and legs is as in 
angelicus. 

Male. — Length 0.8 mm. The single male specimen available for 
study has the antennal pedicel, first funicular segment, and the entire 
club white, the remainder of antenna black. In angelicus the pedicel 
is brownish, the first two segments of the funicle white, and the club 
mostly white but with its base narrowly blackish. Otherwise I can see 
no differences. 

In both angelicus and ahutrvMis the f rontovertex is densely and very 
finely alveolately sculptured, the alveoli very minute and discernible as 
such only under high magnification. This sculpture is very similar to 
tliat found in alvcolatifronx, new species, described beyond, but in that 
species the aveoli are coarser and more distinct, the frontovertex is 
broader and shorter, and the ocelli are arranged in an equilateral 
instead of a distinctly acute triangle. 

Type locality. — Roseland, Va. 

ry;?e.— U.S.N.M. No. 57325. 

Remarks. — The holotype female and one female paratype were col- 
lected by D. W. Clancy, August 1. 1944, at Roseland, Va., on the bark of 
an apple tree infested with mealybugs and bear laboratory No. 745. 
The male allotype and one female paratype are labeled as having been 
reared from Pscudococcus comstocki (Kuwana) collected Octx^ber 18, 
1944, by Clancy at Colesville, Va., and bear laboratory Nos. 855 and 
850, re.spectively. 

». PSEUDAPHYCUS AUSTRIACUS Mercet 

Ptcudnphycus austrincus Mfbct;t, Uev. Kspan. Eiit., vol. 1, p. 13, flR. 2, 1925. 
The type locality is Weyer, Austria. No host is known. 

10. PSEUDAPHYCUS ORIENTALIS Ferriere 

I'sijudaphycus orimtalis Ffkrikke, Bull. Ent. Ros., vol. 28, p. 315, flg. 1, 1987.— liU 
I'yAjj-jr, Trans. Roy. Ent. Soc. London, vol. 0.3, p. 82, 1043. 

This species was originally de.scribed as a parasite of Pseudococcus 
lilacinus (Cockerell) collected at Los Banos, Luzon, Philippine 
Islands. Le Pellcy gives an account of its habits and biology. 

11. PSEUDAPHYCUS MALINUS, new ipecifi 

Aphvcvs Kp. Hakishi.kr and Clancy, Jotirn. Eoon. Ent., vol. 37, p. WH, 1944. 

The description of orientalis Ferriere fits this species in many par- 
ticulars but differs in respect to the ocellar triangle and the speculum 
of the wing. In ornentali-s the ocellar triangle is said to be acute, "the 
hind ocelli half as far from each other as from tlie front ocellus," and 
the wing is said to have "the hairle.ss streak interrupted below the 



318 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 9» 

middle." In malinics the ocellar triangle is somewhat acute, but the 
posterior ocelli are only slightly closer to each other than to the front 
ocellus and the hairless streak is not interrupted. One inconspicuous 
but seemingly constant character distinguishing malinus from all 
other species examined is that the clypeus has a very narrow brownish 
or blackish anterior border which constitutes a nearly straight and 
continuous dark line between the bases of the mandibles. 

Female. — Length 0.9 mm. Frontovertex, mesoscutum, axillae, and 
scutellum dirty orange-yellow ; temples, cheeks, and face below f rons 
whitish except for a very narrow brownish or blackish line along the 
oral margin between bases of mandibles ; occiput above the neck more 
or less blackish ; antennal scape, pedicel, and f unicle dark brownish 
or fuscous, the last funicular segment usually mostly whitish; club 
white ; apices of mandibles dark brown ; eyes black ; ocelli dark red- 
dish ; pronotum whitish above, its anterior face in large part blackish ; 
anterior margin of mesoscutum along the suture narrowly blackish; 
scutellum frequently with indications of an obscure brownish trans- 
verse band before apex; tegulae pale basally, fuscous apically; all 
legs and underside of thorax whitish, the middle and hind tibiae some- 
times each with two more or less distinct fuscous bands ; propodeum 
and dorsum of abdomen dark brown, the latter with a narrow trans- 
verse band of blackish at about basal one-third ; under side of abdomen 
blackish with broad whitish margins. Forewing subhyaline, with a 
faint fuscous cloud behind the stigmal vein but without a hyaline 
band. 

Frontovertex about two and one-half times as long as broad, 
granulosely sculptured; inner orbits parallel; ocelli in a nearly 
equilateral triangle, the posterior ocelli slightly closer to each other 
than to the anterior ocellus; ocellocular line equal to about half the 
diameter of an ocellus; anterior ocellus situated distinctly above 
middle of frontovertex; eyes clothed with short hairs. Antennae 
strongly clavate; scape subcylindrical, approximately four times as 
long as thick; pedicel about one-third as long as scape and subequal 
to first four funicular segments combined; funicular segments all 
transverse and successively increasing in width from first to last, the 
fifth segment fully twice as wide as the first; club solid, distinctly 
wider than last funicular segment and approximately as long as 
pedicel and entire funicle combined. 

Mesoscutum, axillae, and scutellum finely sculptured, mat, and 
clothed with short pale hairs; propodeum very faintly reticulated, 
shining; mesopleuron with very fine and weak reticulation. Forewing 
with short dense discal ciliation distad of speculum, the cilia basad 
of speculum longer and not quite so dense ; speculum closed at posterior 
margin of wing by about three rows of hairs, otherwise not inter- 



CHALCID-FLIES OF GENUS PSEUDAPHYCUS GAHAN 319 

riipted; marjiinal vein piinctiforin; sti*rnial vtMii loiif^^T tliau marginal 
anil postniarginal conibinod. 

Abdomen abont as long and as broad as thorax, short-ovate to nearly 
circular in out lino, weakly reticulated and more or less shining dor- 
sally and vi'ntrally; ovipositor exserted approximately one-sixth to 
one-fifth the length of abdomen. 

Male. — Length 0.75 nnn. Like female in all respects except that 
frontovertex is only about twice as long as broad, anterior ocellus 
only slightly above middle of frontovertex, and ahdonun a little 
shorter and narrower than thorax. 

Type locality. — AVinchester, Va. 

Type.—V.S.^.U. No. 57320. 

Remarks. — The holotype female, allotyi)e, and a largvr number of 
paratypes of both sexes were reared by D. AV. Clancy from Pseudo- 
COCCU.S comstockl (Kuwana) collected August 11, 194->, at Winchester, 
Va., and recorded under Charlottesville, Va., laboratory No. 557. 

This species was introduced from Asia into Virginia and several 
other eastern States in 1941-42 by the Bureau of Entomology and 
Plant Quarantine to combat the Comstock mealybug. Numerous 
specimens of the species reared from P. com-'^tockt collected by R. W. 
Burrell and C. P. Clausen in Japan and Chosen have been examined 
and compared with the types, but since none of this Asiatic material 
is in fir.st-class condition the description has been drawn from speci- 
mens representing recoveries of the species in Virginia. Since its 
original introduction specimens have been distributed for colonization 
in other localities, and the National Collection now contains material 
representing recoveries at the following points: Batesville, Berryville, 
and Hamilton, Va.; Proctorville, Ohio; South River, Morganville, 
:in<l Moorestown, N. J.; Branfurd, Guilford, and Meriden, Conn.; 
;iiid Bridgeville, Del. 

12. PSEUDAPHYCUS ANGUSTIFRONS. new speoiia 

Thf- unusually narrow frontovertex will distinguish tJiis species from 
all others known to me. 

Female. — Length 0.8 mm. Fi-ontovcrtex light orange; occiput more 
or less blaekish above the neck: eyes black; ocelli dark red: remainder 
of head whitish. Antennal scape and pedicel black or bhickish, the 
scape usually pale at base and .sometimes with the dor.sal margin j)ale; 
first three segments of funicle more or less fuscous, fourth and (iflh 
M'gments white; club white. Pronotum whitish dorsally and laterally 
but with anterior face mostly blackish; mesoscutinn, axillae, and 
scutellun) brownish orange, with the sutures darker; propodeum dark 
brown; pleura, .sterna, and all legs whitish; abdomen doisally dark 
blown with whitish margins, ventrally entirely while. Wings uni- 
formly faintly fuscous; marginal vein daik fuscous, other veins \m\W\\ 
TooOei 4»; —2 



320 PROCEEDINGS OF THE NATIONAL MUSEUM vol. oe 

Frontovertex unusually narrow, fully three times as long as broad, 
gi'anulosely punctate ; inner orbits parallel or very nearly so ; ocellar 
triangle strongly acute, posterior ocelli separated by a distance equal 
to r.bout one to one and one-half times the diameter of an ocellus, 
nearly twice as far from anterior ocellus as from each other, almost 
touching the eye margins and a little more than the diameter of an 
ocellus from the occipital margin; anterior ocellus situated a little 
above the middle of frontovertex; eyes large, with very short and 
inconspicuous pile. Antennae strongly clavate ; scape distinctly a 1 ittle 
exj^anded beneath, widest beyond middle, very slightly less than four 
times as long as broad; pedicel about one-third the length of scape 
and about equal to first four segments of f unicle combined ; funicular 
segments all wider than long, successively increasing in width from 
first to last, the first segment narrower than pedicel, the fifth segment 
the largest and about twice as broad as long; club solid, distinctly 
broader than last funicular segment, and approximately as long as 
pedicel and funicle combined. 

Mesoscutum, scutellum, and axillae finely sculptured but not wholly 
mat, clothed with short whitish hairs ; propodeum smooth and shining; 
mesopleuron weakly sculptured. Forewing with speculum closed at 
posterior margin of wing and also interrupted a little behind tlie 
middle ; discal ciliation distad of speculum short and moderately dense, 
proximad of speculum a little coarser and not quite so dense ; marginal 
vein punctiform, postmarginal a little longer than marginal, stigmal 
distinctly longer than marginal and postmarginal combined. 

Abdomen as broad as thorax and usually a little longer than thorax, 
ovate, weakly sculptured dorsally, smooth and shining ventraily; 
ovipositor sheaths exserted about half the length of abdomen. 

Male. — Length 0.7 mm. Closely resembling female but with fronto- 
vertex only slightly more than twice as long as broad ; anterior ocellus 
located very nearly at middle of frontovertex; abdomen no longer 
than thorax and subcircular in outline. 

In some specimens of both sexes the prescutum is narrowly bordered 
anteriorly with black and in some males the ocellar triangle is blackish. 

Type locality. — Cuba. 

Ty/^e.— U.S.N.M. No. 57327. 

Remarks. — Described from 26 females and 4 males, all intercepted 
at quarantine, at three different ports of entry, on shipments of pine- 
apples originating in Cuba and in each case infested with mealybugs 
(probably Pseudococcus hrevipes Cockerell). In most instances the 
parasites were actually reared from the mealybugs. The holotype 
female and two female paratypes were reared at Detroit, Mich., April 
18, 1932, by W. W. Wood, under Detroit No. 1094. The allotype and 
three female paratypes were reared April 28, 1936, at New York, and 



CHALCID-FLIKS OF GLNUS PSEUDAPHYCUS GAHAN 321 

bear New York No. 58589. Other paratypes were intercepted May 
13, 1D3G, uiKler New York No. 59153; Doceniber 28, 19;}L>, under New 
Orleans No. 5953; June 3, 1934, under New Orleans No. 8715; April 
>20, 193G, under New Orleans No. 17;V20; May 22, 1934. under New 
Orleans No. 8-lSS; and April 28, 193r), under New York No. 58577. 

18. PSEUDAPHYCUS MUNDUS, new species 

I'seudococcobiun terryi (FuUaway) Bynum (not tenyi Fullaway), Jdui'h. Ec«>u. 

Eiit., vol. 30, pp. 75G-7G1, 1937. 
I'lciidaphynts sp. nov. Wolcott, Insectae Borlnquenses, pp. 128, .''i2!), 103U. 

This species closely resembles angeUcus (Howard) but may be dis- 
tinguished from that species by the fact that the forewing is more 
weakly and more uniformly infuscated, without a transverse hyaline 
bund, and the speculum is not interrupted. The female differs further 
from angdicU'S by having the antennal club black or blackish at base 
instead of entirely white. The male differs from the male of angelicus 
by having the entire funicle of the antenna nearly uniformly pale 
grayish instead of mostly black with the first two segments only pale. 

Female. — Length 1.1 mm. Frontovcrtex, mesoscutum, axillae, and 
scutellura dilute orange with a slight dusky tinge ; head except fronto- 
vcrtex, tegulae, pronotum, entire underside of thorax and all legs 
white or whitish; propodeum and dorsum of abdomen blackish, 
abdomen beneath whitish; ovipositor pale at base, black at apex; 
eyes brownish black, ocelli dark reddish; occiput immediately above 
tlie neck more or less infuscated and suture between pronotum and 
mesoscutum often fuscous; antennal scape black with a white dorsal 
margin; pedicel, entire funicle, and usually the greater part of basal 
segment of club black; last two segments and apex of first segment of 
club pure while; wings uniformly faintly fuscous, almost hyaline. 
Ilairs on mesonotum white. 

Frontovcrtex two and one-half to three times as long as bioad, 
granulosely punctate; inner orbits practically parallel; ocelli in a 
slightly acute triangle; anterior ocellus situated distinctly above the 
middle of frontovcrtex, lateral ocellus about its own diameti-r from eye 
margin; eyes clothed with short pile. Antenna strongly clavale; 
scape distinctly a little expanded beneath, about four times as long as 
bnjad ; pedicel a little more than one-third as long as sctipe and nearly 
equal in lenglii to four following segments of funicle; funicular seg- 
ments all strongly tiansverse, sube(]ual in length but successively in- 
cieasing in width, the fifth segment approximately twice as wide as the 
first and three times as broad as long; club large, indistinctly 3-seg- 
inented, bn^ader tlian last funicular segment, and as long as or a little 
longer than entire funicle. 

Mesoscutum, axillae, and scutellum weakly .sculptured, rather dis- 
tinctly shining and sparsely haii'y; propodeum smooth: mesopleuron 



322 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

uniformly weakly reticulated. Forewing Avith discal cilia basad of 
speculum a little sparser and slightly longer than distad, the speculum 
closed at posterior margin of wing by several rows of hairs but other- 
wise uninterrupted; marginal vein a little longer than broad, post- 
marginal distinct, about as long as stigmal. 

Abdomen as long as head and thorax combined, as broad as thorax 
at base and tapering to a point at apex, distinctly reticulately sculp- 
tured over the whole dorsal surface, nearly smooth ventrally. Oviposi- 
tor exserted one-third the length of abdomen. 

Male. — Length 0.85 mm. Like female except as follows: Whole 
antenna much paler in color, the scape whitish with a longitudinal 
fuscous stripe, pedicel and funicle grayish white, and club white; 
abdomen not longer than thorax, broadly rounded at apex, nearly 
circular, blackish dorsally but with lateral margins broadly margined 
with whitish. 

7'ype locality. — Houma, La. 

Type.—\J. S. N. M. No. 57328. 

Remarks. — Described from numerous specimens received from D. W. 
Clancy of the Charlottesville, Va., laboratory of the Bureau of Ento- 
mology and Plant Quarantine, where they were reared from Pseudo- 
coccus honinsis (Kuwana) collected at Houma, La., in November 
1943. Besides this type series the following identifications of this 
species have been made by me : One specimen labeled as having been 
reared February 10, 1916, from Pseudococcus calceolariae (Maskell) 
[ — misidentification of P. honinsis (Kuwana)] collected at Audubon 
Park, New Orleans, La., by E. R. Barber ; three specimens reared in 
1943 from Pseudococcus honinsis (Kuwana), collected at Houma and 
Raceland, La., by E. K. Bynum; two specimens indicated as having 
been reared from P. honinsis at Cairo, Ga., in 1932, by E, K. Bynum ; 
two specimens taken at quarantine at Brownsville, Tex., on cut flowers 
from Mexico; nine specimens reared from the pink mealybug of 
sugarcane {THonymus sacchari Cockerell), December 28, 1932, at Rio 
Piedras, Puerto Rico, by F. Sein, under P. R. Ace. No. 178-32. Also 
before me are specimens reared in breeding tests at the Charlottesville, 
Va., laboratory from the following hosts : Phenacoccus gossypn Town- 
send and Cockerell, Pseudococcus comstocki (Kuwana), and P. adoni- 
dvm (Linnaeus). 

This species has become confused in the literature with Aphycus 
terryi FuUaway {^Pseudococcohius terryi FuUaway). In the arti- 
cle by E. K. Bynum cited above, it is indicated that living material 
of A. terryi was received in 1932 by T. E. Holloway, of the Bureau of 
Entomology and Plant Quarantine laboratory in New Orleans, from 
the Hawaiian Sugar Planters' Experiment Station in Hawaii. The 
original stock was apparently increased by propagation in the labora- 



CHALCID-FLIES OF GENUS PSEUDAPHYCUS — GAHAN 323 

tory. and during the same year releases are said to have been made 
on three sugtircane plantations in Louisiana; at Cairo, Ga.; at Belle 
Glade. Fla.; and a small shipment sent to G. N. Wolcott for release 
in Puerto Rico. Other releases of the parasite are said to have been 
made in 1933, 1934, and 1936 in several additional localities in the 
three states mentioned. According to Bynum the parasite was re- 
covered in the field of introduction at Houma in 1932 and each year 
thereafter up to the time of publication. Recoveries were allegedly 
made at several other points of introduction including Cairo, Ga., and 
Belle Glade, Fla. At some points of introduction no recoveries were 
made. According to Wolcott attempts in Puerto Rico to recover 
terryi resulted only in the rearing from Trionymus saccJmri of speci- 
mens which were identified b}- Muesebeck as a new species of the genus 
Pseiidaphycus. 

In 194:3 I received from D. W. Clancy, of the Charlottesville, Va., 
laboratory, a series of specimens of a parasite reared from Fseudococ- 
OU3 honirusis that had been sent to him by J. W. Ingram, of the Bureau 
of Entomology and Plant Quarantine laboratory at Houma, La. 
These specimens had been reared from mealybugs collected in 1943 
from fields in the vicinity of Houma and were supposed to be repre- 
sentatives of the introduced Aphycus terryi. They proved not to be 
A. terryi^ however, but the above-described new species, Pseudaphycus 
inundus. 

Following this discovery a request was made to the Houma labora- 
tory for samples of the original shipment of parasites received from 
Hawaii for introduction into Louisiana. Such a sample was received 
and proved to be true A. terryi. Another lot labeled, "Cairo, Ga., 
1932, ex Psev/lococcufs honinsis E. K. Bynum SC #333," and appar- 
ently constituting a part of the original release at that point, was also 
received. This sample proved to be not terryi but Pseudaphycus 
mundus. No representatives of the other releases were obtained, but 
.specimens taken at Houma and Ilaceland, La., in 1943, supposedly 
representing recoveries of terryi^ again turned out to be P. mundwi. 
The specimens mentioned by Wolcott as having been taken in Puerto 
Rico and identified by Muesebeck as Pseudapkycus n. sp. were located 
in the National Museum collection and were also found to be the same 
as those from Louisiana and Georgia. 

Tlie above data show that the parasite received from Hawaii was 
certainly Aphycus terryi but that this species has not been recovered at 
any of those points of release frcjni which material has been submitted 
for identification. All the alleged terryi reared from field a)llect ions 
at Houma that I have seen have been the new species Pseudaphycus 
munduji. Api)arently alsf) it was this sj)ocies, not terryi., which was 
released at Cairo, Ga. Furthermore, it ap[)ears probable that this was 



324 PROCEEDINGS OF THE NATIONAL MUSEUM vod. oe 

the species introduced into Puerto Rico and that this introduction was 
successful even though the species introduced was not the one intended. 
Pseudaphycus mundus may be indigenous in Louisiana. As noted 
above it was reared at Audubon Park, New Orleans, as early as 1916, at 
least 16 years prior to the attempted introduction of Aphycus terryi. 
Circumstantial evidence would seem to indicate that in the attempt to 
increase the stock of the Hawaiian parasite in the laboratory, field- 
collected material of the host which had already been attacked by 
P. mundiis was introduced into the cages and the two species thus 
became confused. 

14. PSEUDAPHYCUS LDIATULUS, new Bpceies 

The dorsum of the thorax in this species is more weakly sculptured, 
more distinctly shining, and of a darker yellowish-gray color than in 
any of the other species. Most closely resembles graminicola but 
distinguishable by the above characters and its slightly narrower 
frontovertex. 

Female. — ^Length 0.75 mm. Frontovertex light orange-yellow ; occi- 
put slightly fuscous; eyes black; ocelli reddish; remainder of head 
whitish. Antennal scape and pedicel pale yellowish gray; funicle 
black or blackish ; club mostly white, more or less stained with blackish 
basally. Mesoscutura, scutellum, and axillae yellowish gray; prono- 
tum whitish, with its anterior face stained with blackish on each side of 
neck; propodeuni dark brown; pleura, sterna, and all legs whitish; 
tegula whitish, its apex stained with fuscous ; abdomen dorsally mostly 
dark brown or blackish, ventrally varying from mostly blackish to 
mostly whitish with only the middle blackish. Wings nearly uni- 
formly faintly fuscous with faint indication of a paler transverse band 
beyond apex of venation. Ovipositor mostly dark brown, paler at 
base. 

Frontovertex about two and one-half times as long as broad, mat, 
the sculpture made up of extremely minute alveolate punctures ; inner 
orbits parallel for most of their length but diverging rapidly just 
above the scrobe ; ©cellar triangle nearly equilateral, or slightly acute ; 
posterior ocellus about half its own diameter from eye margin and 
about its own diameter from occipital margin; anterior ocellus dis- 
tinctly above middle of frontovertex; eyes with very short, incon- 
spicuous pile. Antennae clavate ; scape not expanded beneath ; pedicel 
approximately one-third as long as scape and about as long as follow- 
ing four segments combined ; funicular segments all broader than long, 
successively increasing in width from first to last, the fifth segment 
about twice as broad as long; club solid, distinctly broader than last 
funicular segment and only slightly shorter than funicle and pedicel 
combined. 



CHALCID-FLIES OF GENUS PSEUDAPHYCUS — GAHAN 325 

Mesoscutuni, axillae, and scutelhim nearly flat, weakly sculptured, 
distinctly shining, and clotlied with short pale pubescence; propo- 
(hnirn smooth and shining: mesopleuron weiikly sculptured. Speculum 
of forewing purtially interrupted near middle and closed at posterior 
margin of wing b}' one or two rows of cilia; discal ciliation moderately 
dense, very slightly coarser basad of speculum than distad of it; mar- 
ginal vein punctiform, postmarginal short, stigmal about twice as 
long as marginal and postmarginal combined. 

Abdomen ovate, very slightly longer than thorax and slightly nar- 
rower than thorax; tergites with distinct, shallow, reticulate sculpture, 
somewhat shining; sternites practically smooth and shining; oviposi- 
tor sheaths exserted approximately one-fifth the length of abdomen. 

Male. — Length 0.G5 nnn. Similar to female but with frontovertex 
only about twice as long as broad, mesoscutum along its anterior mar- 
gin narrowly black, abdomen distinctly shorter and narrower than 
thorax and circular in outline. 

Type locality. — Stevensville, Kent Island, Md. 

ry/>e.— U.S.N.M. No. 57;3i:i). 

Remarks. — Described from 21 females (1 holotype) and 5 males 
(1 allotype) reared by H. S. McConnell, from Phenacoccus sp. found 
feeding on a species of Andropogon at Stevensville on Kent Island in 
Chesapeake Bay, August 3, 1942. 

IS. PSEUDAPHYCUS HERACUS, new species 

This species is very similar to meritoritts, new species, described 
elsewhere in this paper, and the two species are found infesting appar- 
ently the same species of mealybug in some localities in Virginia. 
It differs from jrwritorius, however, by having the segments of the 
funicle shorter and more transverse, by having the club somewhat 
longer than the entire funicle, and by having the anterior ocellus 
situated distinctly above the middle of frontovertex. 

Female. — Length 0.8G mm. Frontovertex, mesoscutum, axillae, 
and scutellum orange with no dark markings; head except frontover- 
tex, tegulae. ])r()notum, pleura, sterni, and all legs whitish; metanotura 
and propodeum orange, a shade lighter than scutellum ; abdomen dor- 
sally mostly yellowish with a narrow blackish cross stripe connecting 
the cerci and basad of this stripe with one and apicad of it with two 
similar but less distinct fuscous stripes; abdomen beneath slightly 
paler; ovipositor pale at base, darker apically; eyes grayish black, 
(xpjli dark red ; antennal scape nearly uniformly pale yellowish, pedi- 
cel and funicle fuscous, the apical funicular segments. usually paler 
than the basal ones; club white. Wings uniformly subhyalinc; mar- 
ginal and stigmal veins fuscous. Hairs on mesonotum whitish. 

Frontovertex somewhat more than twice as long sis broad, finely 
granulose but under high magnification with faint indications of 



326 PROCEEDINGS OF THE NATIONAL MUSEUM vol. »« 

alveolate sculpturing ; inner orbits parallel for most of their length ; 
ocelli forming a slightly acute triangle ; anterior ocellus about one and 
a half times the width of f rontovertex above the edge of scrobal cavity ; 
lateral ocellus less than its own diameter from eye margin and more 
than its diameter from occipital margin ; eyes with, distinct short pile. 
Antenna clavate; scape not expanded beneath, slightly broader at 
middle than at either end, four times as long as broad ; pedicel about 
one-third as long as scape, very nearly as long as first four funicular 
segments combined; funicular segments all broader than long, suc- 
cessively increasing in thickness from first to last, the basal segments 
distinctly narrower than pedicel, last segment about as broad as 
pedicel; club large, broader than last funicular segment and a little 
longer than entire funicle. 

Mesoscutum, axillae, and scutellum uniformly granulosely punctate, 
mat; propodeum smooth; mesopleuron distinctly finely reticulate- 
punctate, subopaque. Forewing with rather dense discal ciliation, 
the cilia basad of speculum a little coarser than those beyond ; speculum 
closed posteriorly by one or two rows of hairs, otherwise uninter- 
rupted; marginal vein punctiform; stigmal vein a little longer than 
postmarginal. 

Abdomen short ovate, slightly narrower and a little shorter than 
thorax, very weakly reticulated and shining dorsally, and practically 
smooth ventrally. Ovipositor exserted about one-fourth the length 
of abdomen. 

Male. — Length 0.75 mm. Similar in practically every respect to the 
female except that the abdomen is nearly circular in outline, obviously 
shorter and narrower than the thorax and blackish medially with the 
margins broadly yellowish. 

T'i/pe locality. — Hamilton, Va. 

Type.—V.S.'^.M. No. 57330. 

Remarks. — Holotype female, allotype, and 13 paratypes reared July 
5, 1943, from Pseudococcus virgatus (Cockerell) , collected at Hamilton, 
Va., and received from D. W. Clancy of the Charlottesville, Va., lab- 
oratory under No. 628 ; also 35 paratypes reared May 16, 1944, from 
the same host collected at Batesville, Va., under Clancy's No. 611. 
Many additional specimens reared by Clancy from P. virgatus col- 
lected at Hamilton, Batesville, Crozet, Winchester, Greenwood, and 
Covesville, Va., are in the collection identified as this species but not 
considered a part of the type series. 

16, PSEUDAPHYCUS ALVEOLATIFRONS. new species 

Differs from all other forms studied in the coarser and much more 
distinct alveolate sculpture and paler orange-yellow color of the 
frontovertex. 



CHALCID-FLIES OF GENUS PSEUDAPHYCl'S (iAHAN 327 

Female. — I-/en«rth O.S'2 mm. Frontovertox very dilute orange yellow, 
its entire surface uniformly densely covered with fine but perfectly 
distinct facetlike punctures. In all other respects apparently agree- 
ing with the foregoing description of wrrar?As so that the description 
of that species except for the frontovertcx will serve for this species 
also. 

Mall. — The head is gone from the single male specimen received 
so that the characters of the head are unknown. With respect to the 
other characters this male seems to differ in no significant respect from 
the male of meracus. 

Type locality. — North Bergen, N. J. 

Type.—V.S.^.^l. No. 57331. 

/ifmarl.-s. — Described from a unique female and one headless male 
reared in November 1938 by George Rau from Pseudococcus com- 
^tocki (Kuwana). 

The frontovertcx in most of the forms studied is granulosely sculp- 
tured and mat, with little or no indication of distinct separate alveoli 
or punctures even under the higher magnifications of a binocular 
microscope. In anf/eh'cus, ahstru^ii-s, meraou.s., and limafulits the sculp- 
ture of frontovertex is more or less distinctly alveolate but the alveoli 
are finer and less definite. P. alveolatifrons differs from angeliciis and 
ahstnuiiwi further by having the ocelli in an equilateral, instead of a 
distinctly acute, triangle and l)y lacking the transverse hyaline streak 
on the wing. From Umatulus it may be distinguished by the distinctly 
duller sculpturing of the mesonotum as well as by its brighter orange- 
yellow color. 

It is possible that additional material may show that the differences 
between meracus and aJ^vcolatifrons are not of specific value, but in 
view of the fact that the types of the two species are from different 
hosts and from different localities and this sculptural difference does 
Lxist, it seems wisest to give a separate name to this form from 
/-'. comntocki even though it may eventually turn out to be a synonym. 



I tOVlaNMIIIT PRINTJRt OFriCI II4« 



PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 



iutui ISiJlvA, 0*0 hy the 




SMITHSONIAN INSTITUTION 

U. S. NATIONAL MUSEUM 
Vol. % Wa.hington: 1946 No. 3201 



NEW CERAMBYCID BEETLES BELONGING TO THE TRIBE 
DISTENIIXI FROM CENTRAL AND SOUTH AMERICA 



By W. S. Fisher 



DuRixG the process of rearranging the American species of the tribe 
Disteniini (Coleoptera: Cerambycidae) in the United States National 
Museum, four new species were found. These are described herein. 

Genus DISTENIA Lepelitier and ServiUe 
DISTENIA LATERALIS, new species 

Head, pronotum, and underside of body reddish brown; elytra and 
scutollum yellowish brown, the elytra with a distinct, broad, longitudi- 
nal, green vitta on each side near lateral margin ; antennae brownish 
black; palpi and legs palo brownish yellow. 

Head glabrous, sparsely, irregularly punctate. Antenna nearly 
one and one-half times as long as body, sparsely clothed with long 
flying hairs on underside: first segment slightly shorter than third, 
slender, cylindrical, gradually expanded toward apex, not depressed 
on top at base, rather densely, finely punctate, and sparsely clothed 
with long and short, semierect hairs. 

Pronotum, not including lateral spines, slightly longer than wide, 
widest at middle; sides strongly constricted near base iind ajiox, tri- 
angidnrly expanded on each side at middlo jind ai-med with a long, 
f^onical, acute s[)ine; disk broadly, transvei-sely depressed near autcfior 
margin, transversely, sinuately grooved near base, with a sliglitly 
ol'vatod space oji each side in front of transvoi-se, basal groove; sur- 
face glabrous, coarsely, sparsely, irregularly puiu-tate. with a smooth, 
elongated, median space. 

Elytra at base slightly wider than pronotum including lateral 
spines; sides gradually converging from bases to apices, wliieh are 
T13015— «« 320 



330 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 9fl 

unispinose, the lateral spine on each rather short and acute, and the 
sutural angle feebly produced; surface very densely, coarsely, deeply 
punctate, becoming nearly impunctate near apices, very sparsely 
clothed with rather short, inconspicuous, erect, yellowish hairs. 

Body beneath indistinctly punctate, very sparsely, irregularly 
clothed with short, inconspicuous, recumbent, yellowish hairs, with a 
few longer, erect hairs intermixed; femora slender, cylindrical, 
slightly clavate, each armed with a long, acute spine at apex. 

Length 22 mm., width at base of elytra 4.5 mm. 

Type locality. — Rurrenabaque (Beni River), Bolivia. 

T^^z/^e.— U.S.N.M. No. 57612. 

Remarks. — Described from a single specimen collected at the type 
locality during October by William M. Mann in connection with the 
Mulford Biological Exploration during 1921-1922. 

This species is closely allied to Distenia limhafa Bates, but it differs 
from that species in having the head and pronotum reddish brown 
with only a vague bronzy reflection, the lateral spines on each side of 
the pronotum long and acute, the scutellum yellowish brown, the elytra 
more coarsely and densely punctured and with the apices armed with 
a long, acute, lateral spine, and the antennae brownish black. 

DISTENIA SPINIPENNIS. new species 

Head, pronotum, scutellum, elytra, and underside of body greenish 
black (elytra slightly more greenish) ; antenna (except first segment 
which is black) and palpi reddish brown; legs pale brownish yellow, 
the femora slightly darker at apices. 

Head nearly glabrous, sparsely, irregularl}', indistinctly punctate. 
Antenna one and one-half times as long as body; first segment dis- 
tinctly shorter than third, robust, very strongly clavate, strongly, 
longitudinally depressed on top on basal half, scabrous and coarsely 
punctate basally, and sparsely clothed with long, fine, semierect hairs. 

Pronotum, not including lateral spines, slightly longer than wide, 
widest at middle ; sides strongly constricted near base and apex, tri- 
angularly expanded on each side at middle and armed with a rather 
long, acute spine; disk broadly, transversely depressed near anterior 
margin, narrowly, transversely, sinuately grooved near base, narrowly, 
transversely grooved along base, and with three slightly elevated, 
elongate, smooth spaces, one median and one on each side of middle; 
surface sparsely, indistinctly punctate in depressions, and clothed with 
a few short, indistinct, erect hairs. 

Elytra at base as wide as pronotum including lateral spines ; sides 
strongly converging from bases to apices, which are unispinose, the 
lateral spine on each long and acute, and the sutural angle rectangu- 
lar; surface coarsely, densely, deeply punctate, becoming nearly im- 



NEW CERAMBYCID BEETLES — FISHER 331 

punctate at apices, rather densely, uniformly clothed with short, erect, 
whitish hairs. 

Body beneath indistinctly, irre«;ularly punctate, sparsely clothed 
toward sides with long and short, seniierect, whitish hairs; femora 
slender, slightly clavate, unarmed at apices. 

Length 19.5 mm., width at base of elytra 4 mm. 

Type lotality. — "Guapiles," 250-300 meters, Costa Rica. 

Ty/^d.— U.S.N.M. No. 57613. 

Remarks. — Described from a single specimen collected at light dur- 
ing May 1934, by Ferdinand Nevermann. The specimen is labeled 
Guapiles, which is probably an error for Guaplies. 

This species is closely allied to Distenia phaeocera Bates, but it 
dili'ei-s from the description given for that species in not having the 
elytra stria-punctate or the surface alutaceous, and in the slightly 
elevated smooth spaces on each side of the pronotum being not divided. 

Genus COMETES Lepelitier and Serville 

COMETES EMARGINATA. new species 

Elongate and strongly shining, black, except anterior coxae, anterior 
femora (except tips) , basal halves of middle and posterior femora, and 
tibiae in part brownish yellow. 

Head finely, irregularly punctate on top, somewhat transversely 
rugose anteriorly, with a smooth, longitudinal groove extending from 
clypt'us to occiput, sparsely clothed with long, erect and recumbent, 
whiti.-h hairs. Antenna one and one-half times as long as body, 
sparsely clothed with l<->ng, flying hairs on underside; first segment 
slightly longer than third, robust, cylindrical, gradually expanded 
toward apex, finely, rugosely punctate, sparsely clothed with long erect 
and short recumbent, whitish hairs. 

Pronotum as wide as long, widest at middle; sides strongly con- 
stricted near base and apex, triangularly expanded on each side at 
middle but not distinctly spinose; disk broadly, transversely depressed 
near anterior margin, transversely, sinuately grooved near base, nar- 
rowly, transversely grooved along ba.se, and with two slightly elevated, 
rouiul, smocjth gibbosities on each side <jf middle; surface densely, 
finely punctate; in median depression, the punctures nearly concealed 
by a dense, recumbent, yellowish pubescence, and with a few long, erect, 
white hairs. 

Elytra at base as wide as [)ron<jlum at middle; sides gradually con- 
verging from bases to apices, which are separately enuirginale. the 
lateral spine slightly longer than su(ural si)ine; surface coarsely, 
deeply, densely punctate, tiie puncture forming more or le.ss distinct 
rows but becoming obsolete toward apices, rather den.sely clothed 
toward sides with short, indistinct, it'iinibent, whitish hairs, with a 



332 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96 

few long, erect liairs intermixed, and each elytron ornamented with a 
narrow vitta of transversely recumbent, white hairs along sutural 
margin. 

Body beneath indistinctly punctate, sparsely clothed at sides with 
short, recumbent, whitish hairs ; legs sparsely clothed with long and 
short, semierect, white hairs. 

Length 11 mm., width 2 mm. 

Ti/pe locality. — Hamburg Farm (on Keventazon River) , Costa Rica. 

Type.—V.S.'NM. No. 57614. 

Remarks. — Described from a single specimen collected at the type 
locality on grass, May 26, 1934, by Ferdinand Nevermann. 

This species can be separated from the other described species of 
this genus by having the tips of the elytra emarginate. 

COMETES BICOLOR, new species 

Elongate, rather strongly shining except elytra which are sub- 
opaque; antennae bluish black with a vague violaceous reflection; 
head, pronotum, scutellum, underside of body, and legs bluish black 
with a distinct greenish or violaceous tinge; elytra brownish yellow 
with apical fourth and elevated lateral margins violaceous blue. 

Head rather coarsely, irregularly punctate on top, somewhat trans- 
versely rugose anteriorly, with a smooth, longitudinal groove extend- 
ing from clypeus to occiput, very sparsely clothed with inconspicuous, 
semierect hairs. Antenna nearly one and one-half times as long as 
body, sparsely clothed on underside with long, flying hairs ; first seg- 
ment as long as third, slender, cylindrical, narrow at base, strongly 
expanded toward apex, rather densely, coarsely, shallowly punctate, 
sparsely, uniformly clothed with short, semierect, whitish hairs. 

Pronotum slightly wider than long, widest at middle ; sides strongly 
constricted near base and apex, obtusely expanded on each side at mid- 
dle but not spinose ; disk broadly, transversely depressed near anterior 
margin, shallowly, transversely, sinuately grooved near base, nar- 
rowly, transversely grooved along base, with two irregular, smooth 
elevations on each side of middle, and an elongate, smooth, median 
elevation; surface coarsely, deeply, confluently punctate, sparsely 
clothed with moderately long, erect, inconspicuous hairs. 

Elytra at base as wide as pronotum at middle; sides parallel from 
bases to near apices, which are separately narrowly rounded ; surface 
densely, coarsely, deeply, uniformly punctate, the punctures becom- 
ing more confluent toward apices, rather densely clothed with short, 
inconspicuous, erect hairs, and each elytron with two more or less dis- 
tinct longitudinal costae, the inner costa extending from base to mid- 
dle of elytron, and outer one from base to apical fourth. 

Body beneath sparsely, shallowly, irregularly punctate, sparsely, 
irregularly clothed with moderately long, semierect, fine hairs; legs 
rather densely clothed with long and short, semierect, whitish hairs. 



NEW CERAMBYCID BEETLES — FISHER 333 

Length 11.5 mm., width 3 mm. 

Type locality. — Colombia, South America (no definite locality). 

ry^e.— U.S.X.M. No. 57G15. 

litmarlca. — Described from a single specimen collected August 31, 
1942, by Francisco J. Otoya (No. 2053). 

This species is allied to Comctes pulcherrivins Bates, but it differs 
from that species in iuiving the pronotum more densely and coarsely 
punctured, and the basal three-fourths of the elytra brownish yellow, 
with only the apical fourth and narrow lateral margin of each elytron 
violaceous blue. The head is mounted separately on the same pin 
with the rest of the specimen. 



u I aaviNMHiNr PMiNTiMa orricit lt4t 



PROCEEDINGS OF THE UNITED STATES NATIONAl MUSEUM 




issued jvAjfV vi» Cjyj ^ fy the 

^^^^ 

SMITHSONIAN INSTITUTION 
U. S. NATIONAL MUSEUM 



V' I. ')6 Washingtor: 1946 No. 3202 



MACHAEROIDES EOTHEN MATTHEW, THE SABER- 
TOOTH CREODONT OF THE BRIDGER EOCENE 



By C. Lewis Gazin 



The 1940 Smithsonian Institution expedition to the Bridger 
Basin of Wyoming' had unusual good fortune in securing skeletal 
remains of the rare creodont Machaeroides eothen. The materials, 
consisting of skull, lower jaws, and other skeletal portions of one 
individual, were found by Franklin Pearce in low exposures, 
probably of Bridger "C," immediately to the north of Twin 
Buttes, about 30 miles southwest of Green River, Wyo. Previous 
finds of this remarkable sabertooth form, so far as known, are 
limited to the lower jaw portions described by Matthew ^ in 1909. 

The skull, U.S.N.M. No. 17059, v/as found with the lower jaws 
in a position of articulation, and although essentially complete 
there has been a small amount of transverse crushing and dis- 
tortion so that the riglit side of the skull is higher than the left. 
The remainder of the skeleton includes portions of the vertebral 

lumn, both humeri and femora, the right radius and ulna, an 
incomplete left tibia, and fragments of the pectoral and jielvic 
girdlrs. The feet were not present oxcopt for a carpal and two 
metacarpal bones. 

Matthew readily appreciated the indications in the lower jaw 
portions of Machaeroides eothen of a modification nearly parallel- 
ing that of the machairodont forms among the Fissipedia. In 
the materials he had at hand these modifications were not too 
evident, and R. H. Denison - was inclined to consider the M. 
eothen jaw as resembling Felin and not truly "sabertooth." The 

' Matthew, W. D.. Mem. Amcr. Mus. N*t Hist., vol. 9, pt. 6, pp. 4B2-<63, 1909. 
' n..nl<»on. R. II.. Ann. N«w York Acad. 8cl.. vol. 87. p. 181, 1938. 

335 



336 PROCEEDINGS OF THE NATIONAL MUSEUM VOL.96 

National Museum specimen, however, leaves no doubt of the 
direction and extent of specialization, which quite parallels that 
seen in the machairodonts but not to the extent exhibited in the 
Uinta Apataelurus kayi described by Scott. ^ Structurally no 
characters are observed in the illustrations of the lower jaws of 
A. kayi that would preclude derivation of this form from M, 
eothen, with the characteristics attendant upon sabertooth de- 
velopment reaching a high degree of specialization within an 
interval of time suggesting a comparatively rapid evolutionary 
tempo. 

Matthew regarded Machaeroides eothen as an oxyaenid type 
of creodont, closely related to forms included in the subfamily 
Limnocyoninae, to which he allocated it (p. 410) , * having created, 
however, the subfamily name "Machairoidinae" on an earlier 
page (330). Denison ^ retained the supergeneric separation, 
which was entirely justified by the discovery of Apataelurus, 
recognizing a natural phyletic subfamily. However, Denison 
removed the Limnocyoninae and Machaeroidinae from the Oxyae- 
nidae and placed them in the Hyaenodontidae. Justification for 
this was claimed on the basis of morphological differences between 
the Oxyaeninae and Limnocyoninae and similarities between the 
Limnocyoninae and Proviverrinae, particularly between Prolim- 
nocyon and Sinopa. Separation from the Oxyaenidae seems sup- 
ported, and, moreover, carnassial specialization, which has been 
the key to their supposed affinity, may well have developed inde- 
pendently in the two groups. I cannot, however, but regard the 
carnassial differentiation that so readily distinguishes the Lim- 
nocyoninae from members of the Hyaenodontidae as being of 
fundamental importance, and hold that the morphological simi- 
larities may be as easily attributed to similar adaptation or 
parallelism. It is in a similar manner that the Machaeroidinae 
so markedly resem.ble the machairodonts, although obviously not 
derived one from the other. Derivation of the Limnocyoninae 
from an early proviverrine stock is not disproved and may be 
reasonable as suggested by the similarity of lower jaws belonging 
to forms of Prolimnocyon and Sinopa morclax, but their di- 
vergence must originate in a primitive form with molars as yet 
undifferentiated as to carnassials, and where M-, for example, has 
not taken on characteristics so markedly different in the two 
groups. 



3 Scott. W. B., Ann. Carnegie Mus., vol. 27, art. 6, pp. 113-120, 1938. 

* Matthew, W. D., ibid. 

» Denison, R. H., ibid., p. 181. 



MACHAEROIDES EOTHEN MATTHEW GAZIN 337 

Inasmuch as it seems advisable to exclude the Limnocyoninae 
from the Oxj-'aenidae, and since I am unable to reconcile it with 
the Hyaenodontidae on the basis of carnassial differentiation, I 
propose, at the risk of censure, that the Limnocyoninae and 
Machaeroidinae be given family recognition as the Limnocyonidae, 

Description of skull (pi. 45) . — The skull of Machaeroidcs eothen 
is significantly smaller than that of Limnocyori verns but much 
larger than Thinocyon vclox, corresponding closely in size to 
Siyiopa rapax among its less closely related contemporaries. The 
skull is moderately slender but with a noticeably deep rostrum, 
high srgittal crest, and a narrow occiput. 

The rostrum, in addition to depth, exhibits a well-inflated 
maxillary portion covering the long root section of the canine, 
extending parallel and immediately posterior to the suture join- 
ing the premaxilla. The nasals extend posteriorly in a nearly 
F-shaped wedge between the frontals, terminating fully as far 
back as the postorbital processes. In Thinocyon and Limnocyon 
the nasals as exposed terminate distinctly forward of this position. 
The postorbital processes of the frontals are moderately well 
developed and, although possibly not complete, do not appear so 
prolonged as in Thinocyon. The lachrymal bone in M. eothen is 
lar;;e and extends well forward of the orbit and exhibits a most 
unusual feature in that the lachrymal foramen enters the skull 
anterior to the orbital rim and anteroventral to the lachrymal 
crest or tubercle. A smaller foramen also enters the lachrymal 
bone outside and immediately dorsal to the lachrymal tubercle. 
This arrangement was not ob.-.erved in other creodonts, but it 
occurs in marsupials and sloths. Matthew, " however, noted that 
the lachrymal foramen in Limnocyon was very near the orbital 
rim. 

The cranial portion of the skull is characterized by an exceed- 
ingly small brain case, relatively much smaller than in Thinocyon, 
and a very high sagittal crest, which joins a high but narrow 
occipital crest. The latter is overhanging but, quite unlike Lim- 
nocyon, is noticeably constricted transversely immediately above 
the occipital condyles. A prominent vascular foramen is noted 
at the suture between the parietal and squamosal at a position 
about over the trough for the audital tube, and one also in the 
posterior portion of the temporal fossa, presumably at about the 
juncture of the parieto-squamosal and parieto-occipital sutures. 

In ventral aspect the palatal portion of the skull shows no 
unusual features. No evidence exists of the grooves and ridges 



• Matthew. W. D , ibid., p. 438. 



338 PROCEEDINGS OF THE NATIONAL MUSEUM VOL.96 

characterizing' the later fissiped sabertooth cats. However, palatal 
excavations between the deuterocone or protocone portions of 
successive 3-rooted teeth, for reception of the much-elevated pro- 
toconids of the lov/er cheek teeth, are as well developed as in 
several of the other creodonts. The posterior margin of the palate 
is extended posteriorly a very short distance below the narial 
passage, somewhat as in Limnocyon. The palatal margin outside 
. the narial passage, however, shov/s a conspicuous notch for the 
palatine vein, with well-developed tuberosities on both the palatal 
and maxillary sides of the groove. 

^ The zygomatic arch arises from perhaps a slightly higher posi- 
tion on the relatively deep rostrum of Machaeroides eotken than 
it does in Limnocyon verus, although in M. eothen the left arch- 
is broken down to a position below normal on that side of the skull. 
The depth of the arch is moderate, but because of crushing the 
extent to vv^hich it is expanded laterally cannot be determined. 
Posteriorly the arch terminates at a position relatively low with 
respect to the basicranial surface. The zygomatic process of the 
squamosal projects conspicuously dovv^nward from the basicran- 
ium, placing the glenoid surface for articulation of the lower jaw 
at a much lower level than observed in other creodonts, a condition 
noted in machalrodonts and in the sabertooth marsupial Thyla- 
cosmilus atrfT. This lowering of the fulcrum gives leverage to 
the temporal muscle acting on a coronoid process of reduced 
height. The reduction of the coronoid presumably permitted the 
lov/er jaw to open v»'ide enough for the mandible to clear the 
saberteeth, apparently much wider than necessary in other carni- 
vores, except Apataelurus and the machalrodonts. 

The basicranial portion of the skull is the least distorted by 
crushing and is relatively elongate and clearly much narrower 
than in Limnocyon verus. The paroccipital process shows very 
little development and does not project posteriorly as in Limno- 
cyoii verus. However, in addition to the downward-projecting 
pedestallike base for the glenoid surface, the mastoid process, as 
in the machairodonts, is very well developed, projecting downward 
and forvv^ard, and noticeably expanded in an anterointernal- 
posteroexternal direction. The mastoid process is moderately 
developed in most creodonts including Limnocyon, but nowhere in 
the suborder is it relatively so important, particularly in a for- 
ward medial extension, as in M. eithen, and in Apataelurus by 
inference. Its prominence and rugosity demonstrate the strength 
and importance of the sterno-cleido-mastoid muscle, the actions 
of which include depressing the head, as in striking with the 



MACHAEROIPES EOTHKN MATTHEW — GAZIN -^39 

sabers. The mastoid process is essentially a part of the mastoid 
portion of the periotic ; however, the extent to whicli the squamosal 
enters into its composition cannot be determined, limited possibly 
to a portion of the anterior surface of the process. 

The foramina of the hasicranium show certain significant dif- 
ferences in relative position from those in Limiiocyon vems or in 
Thinocyon relox. For the most part these maintain a primitive, 
creodont arrangement with differences noted in Machaeroides 
eothcn that are in part due to structural modification attendant 
upon sabertooth development. The alisphenoid canal, if present, 
is decidedly long as in Limnocyon, but with the posterior opening 
confluent with or not distinguished from the foramen ovale, so 
that it was not certainly identified in the material at hand. In 
Limnocyon veirus the posterior opening of the alisphenoid canal 
's shown by Matthew ^ as well forward of the foramen ovale. 
The foramen ovale in M. eothcn is located medial to the post- 
glenoid process, at the root of the pterygoid wing of the alisphe- 
noid. The postglenoid foramen enters the skull at the base of 
the posterior surface of the postglenoid process but exhibits a 
groove for about half the length of this surface before closure is 
complete. The foramen lacerum medius is in a customary posi- 
tion antero-internal to the exposed portion of the petrosal. From 
a position medial to the promontorium of the petrosal a narrow 
cleft extends posteriorly along the outwardly convex lateral 
margin of the basioccipital to the foramen lacerum posterius, 
about halfway to the occipital condyles. The internal carotid 
evidently entered the brain case at some point along the anterior 
part of the cleft and adjacent to the petrosal. The condylar or 
hypoglossal foramen is well forward of the condyles and sepa- 
rated by a thin partition from the foramen lacerum posterius, 
quite unlike Limnocyon or Thinocyon but resembling certain 
specimens of the Oligocene Daphoeims in this respect. The 
stylomastoid foramen shows as a groove on the medial margin 
of the mastoid process. Dorsally this is completely enclosed, 
presumably by bone belonging to the mastoid portion of the 
periotic, at the root of the mastoid process, posterolateral and 
very close to the promontorium of the petrous portion. 

Machaeroides eothcn is without a tympanic bulla, and the 
tympanic ring was not preserved. The site of the audita] 
tube is a deep and compressed J7-shaped trough between the 
postglenoid and mastoid jjrocesses, extending laterally and some- 
what posteriorly from below the anteroexternal portion of the 

' M«tth-w. W. D.. ibid.. t\z. 66. 



340 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. % 

petrosal. In Limnocyon and Thinocyon the trough is more widely 
open and relatively shorter. The petrosal is partially exposed 
on both sides of the skull of M. eothen. It exhibits an acutely 
projecting promontorium located immediately inward from the 
medial margin of the mastoid process. The petrosal is more 
broadly rounded ventrally in Thinocyon. The fenestra rotunda 
in M. eothen is large and faces posteriorly and slightly outward 
and downward below the flattened ventral surface of the pos- 
terior portion of the petrosal. The anterior surface of the 
petrosal is broad and flattened, facing anterolaterally and ven- 
trally and joining the medial surface in a bluntly rounded angle 
that extends anteromedially and dorsally from the promontorium. 
The fenestra ovale faces slightly forward of lateral and is situ- 
ated very deep in the mesotympanic fossa, almost directly above 
but well separated from the promontorium. It is much higher 
in position than the fenestra rotunda and is very much above the 
lip of the trough for the audital tube. Further description of 
the petrosal and its relation to the cranial cavity is not under- 
taken, inasmuch as such additional information cannot be ob- 
tained without damage to the skull. 

Upper dentition. — The dental formula for the upper teeth of 
Maclmeroides eothen is 3-1-4-2, as noted by Matthew in Lim- 
nocyon and as observed in Thinocyon. Moreover, the first upper 
molar is the carnassial as in these forms and as in the Oxyae- 
nidae. The teeth resemble those in the Limnocyoninae with 
certain exceptional characteristics which for the most part are 
modifications seemingly accompanying sabertooth development. 

The incisors are slender, recurved, conical teeth adapted for 
piercing and with transversely fiattened roots. Unlike Lim- 
nocyon they increase in size from first to third, P being much 
more robust and with a root portion about twice as long as in I'. 
P in Limnocyon is much smaller than P and may not be present 
in some individuals, as indicated by Matthew. * In Thinocyon 
the three are subequal and slightly spatulate. 

The enlarged canine is preserved only on the left side and is 
broken away a short distance below the alveolus. It is removed 
from P by a short diastema. The tooth as exposed has no cingu- 
lum, is nearly oval in cross section, and has a long gently curved 
root section, as inferred from the inflated portion of the maxilla. 
In Matthew's ® illustration of Limnocyon verus the upper canine 
appears to have an exposed cingulum and the short crown is 



• Matthew, W. D., ibid., p. 434. 
•Matthew, W. D., ihid., fig. 58. 



MACHAEROIDES EOTHEN MATTHEW — GAZIN 341 

noticeably recurved below this point. The length of the canine 
in M. eothen cannot be determined from the present material; 
however, if this tooth extended as far as the flange of the lower 
jaw it would have had a length of about 3 cm. beyond the alveolus. 
The canine shows no evidence of serrations along the anterior or 
posterior margins. 

The premolars of M. eothen are 1-, 2-, 3-, and 3-rooted, respec- 
tively. F\ preserved only on the right side when found but 
subsequently lost, is a small, simple, conical tooth without an 
accessory cuspule and separated by a diastema from both the 
canine and P-. In both Limnocyon and Thinocyon P^ is 2-rooted 
and exhibits a small posterior cuspule. P- of M. eothen, in addi- 
tion to being 2-rooted, retains a vestige of a posterior cuspule. 
The tooth is relatively much smaller and transversely more 
compressed than in Limnocyon. P% on the other hand, is much 
better developed than in Thinocyon and apparently than in 
Limnocyon. This tooth, preserved only on the left side, has 
a minute parastyle and a rather well developed posterior cusp 
or crest, approximating P* in this respect. The lingual root 
is slender but extends markedly inward from about midway 
of the tooth length and supports a very small deuterocone. 
P* is relatively robust with a prominent parastyle before and a 
trenchant cusp posterior to the large backward sloping primary 
cusp. The deuterocone portion is broad and well defined, extend- 
ing lingually from the midportion of the tooth. The deuterocone 
portion does not project forward as it does in Thinocyon or (to a 
less degree) in Limnocyon. The deuterocone is situated on the 
lingual margin of the talon and is connected with the primary 
cusp by a low crest across the forward part of the talon. A small, 
shallow basin is enclosed between this forward crest and the 
low cingular crest around the posterior margin of the talon. 

The difference in development of the various premolars is 
rather striking in comparison with related forms. P' and P* are 
relatively small and less progressive, probably reduced from an 
earlier but more advanced state, paralleling the machairodonts in 
this respect, as an accompanying factor in sabertooth specializa- 
tion. P\ however, has retained a more advanced stage of devel- 
opment and undoubtedly continued to be a significant and func- 
tional tooth in the later and more advanced Apntaeluntfi, as 
indicated by the well-developed and trenchant P^ with which it 
occludes in the lower jaw of the Uinta form. 

^r, the upper sectional tooth in Machneroides eothen, has taken 
on a very trenchant appearance as compared either with the 



342 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. w 

preceding P* of MacJiaeroides or with M^ in Limnocyon and 
Thinocyon. The talon portion is much reduced and far forward 
in position, the deuterocone being but a very small cusp at the 
anterolingual angle of the tooth. The posterior crest is elongate 
and the greater part of the lingual face of the tooth forms a flat 
shearing blade which, though oblique, is directed more nearly 
longitudinal than in either Limnocyon or Thinocyon. The para- 
cone and metacone of M^ are closely connate, more so than in 
Limnocyon, approximately as in Tritemnodon. The parastyle is 
but weakly developed, being represented only by an enlargement 
of the cinguium anteroexternal to the paracone. 

M^ has a nearly transverse shear and is characterized in dis- 
tinction from that in Limnocyon and Thinocyon in having lost all 
trace of the metacone. Also, the talon is reduced to a very sub- 
dued projection from the paracone, the shear being effected essen- 
tially by the paracone and parastyle, whereas in Limnocyon and 
Thinocyon the talon participates very largely in the shearing 
structure, cutting against the metaconid and occluding with the 
talonid of Mo. The reduction of the talon of M^ in M. eothen 
accompanies the loss of the metaconid and extreme reduction of 
the talonid of the lower carnassial. 

Mandible (pi. 46, a, b) . — The lower jav/ of Machaeroides eothen 
has been briefly described and figured by both Matthew and Deni- 
son, and the relationship to other creodonts discussed at length. 
However, the specimens available to them for study were incom- 
plete, lacking particularly the extremity of the flange, coronoid 
process, and the crowns of the canine and carnassial teeth. Res- 
torations of these parts in drawings were conservative in indicat- 
ing less modification from the limnocyonine pattern than the more 
complete material demonstrates. 

The lower jaw is relatively very deep and transversely slender 
compared to jaws belonging to forms of the Limnocyoninae. Its 
flange projects downward to a greater extent than anticipated but 
not to the extent shown in Apataelurus kayi, and much less than 
in Hoplophoneus. In M. eothen it projects but a little below the 
dorsoventrally elongate symphysis and curves gently outward so 
that the width across the extremities of the flanges is greater than 
at any point above. The deep symphyseal surface has a nearly 
straight, steeply inclined anterior margin, which makes an abrupt 
angle with lower margin. The coronoid process of the lower jaw 
appears truncated and much reduced from the large and fully 
developed coronoid observed in Limnocyon and Tkhiocyon. The 
reduction, nevertheless, has not gone so far as that in Apataelurus. 



MACHAEROIDES EOTHEN MATTHEW — OAZIN 343 

Moreover, the condyle is lower with respect to the tooth row. 
The angle, less widely separated from the condyle, does not curve 
downward from the nearly straight lower margin of the jaw 
but preserves the alignment practically to its extremity, more so 
than in Apataeh/jus. The masseteric fossa is well defined with 
a sharp masseteric crest anterodorsally ; however, the fossa, as 
in Apataelurus and also Pafriofelis, extends farther forward be- 
neath the carnassial than in Limnocyon and Thinocyon. The 
mental foramina are beneath Pj and the posterior root of P-j and 
placed relatively low on the side of the jaw. 

Lower dentition. — The lower teeth are all present, although 
the right canine and Po as well as the left carnassial and median 
incisors are slightly damaged. The lower incisors of Machae- 
roides eothen were most certainly reduced to two. The lateral 
of these is the larger of the two, and both, as in the case of the 
upper incisors, are piercing type structures with transversely 
flattened roots. In lateral view the canine appears moderately 
robust at the alveolar border but is transversely flattened and 
tapers rapidly to a point only a little above the closely adjacent 
incisors. Moreover, it shows a pronounced scar or bevel verti- 
cally along its posterolateral surface, worn through occlusion with 
the superior canines. 

The cheek teeth appear decidedly slender, and the two anterior 
premolars, both of which, however, retain two roots, are of rather 
small size. P3 is about intermediate in size and development be- 
tween P2 and P^. It has a crested talonid, almost as in P4; a 
structure but very feebly expressed in Po. P3 is without a para- 
conid. whereas in P, this cusp is low but distinct. Both P3 and P4 
are more progressive than in Limnocyon and Thinocyon; however, 
in Apataelurus P.-^ has become relatively much reduced in size. 
P^ in both Machaeroides and Apataelurus has become a relatively 
large and functionally significant tooth, actually exceeding Mi 
in size in both forms. 

The two molars possess somewhat more distinctive structural 
characters than the premolars in characterizing Machaeroides. 
In M, the metaconid is but moderately developed and pressed 
close to the protoconid. The talonid is much reduced in size and 
trenchant in character. In Limnocyon and Thinocyon the rela- 
tively wider and more robust Mi has a better-developed metaconid 
and a large, deeply basined talonid. In Apafaelvrvs the trenchant 
heel persists and evidence remains of a metaconid that has not 
been entirely oliliterated by the shearing function imposed on the 
tooth. 



344 PROCEEDINGS OF THE NATIONAL MUSEUM VOL.98 

The camassial, M2, in M. eothen exhibits more noticeable modi- 
fications than Ml toward a sabertooth specialization. The meta- 
conid is lost, except for a slight rugosity on the posterointernal 
margin of the backward sweeping protocone. The talonid is re- 
duced to little more than a vestige, although its expression is 
somewhat better in the specimen figured by Matthew. ^° The 
tooth is almost catlike in appearance, quite unlike that in Limno- 
cyon or Thinocyon where the carnassial is very much like Mj 
except for its greater size and higher trigonid. In Apataeluncs 
the lower carnassial has almost or quite reached the stage ex- 
hibited by Smilodon in its modification for shearing, but still 
preserving a vestige of its trenchant talonid. 

Table 1. — Measurements {in millimeters) of skull, mandible, and dentition 
of Machaeroides eothen, U. S. N. M. No. 17059 

Greatest length of skull from anterior margin of premaxillae to posterior margin 

of occipital condyles a 135 

Distance from anterior margin of premaxillae to posterior narial aperture a 65 

Greatest depth of maxillae above P' a 32 

Width of palate between canine alveoli a 16 

Greatest width of basicranial region across mastoid processes a 48 

Length of upper dentition from I' to M= inclusive a 62 

Length of upper dentition from anterior margin of canine alveolus to M' inclusive. ... a 66 

Width across I' to P inclusive 76 

Length of cheek tooth series, P' to M- inclusive 42 

Length of diastema anterior to P' : posterior to P^ 3.6 ; 2.6 

Length of premolar series, P^ to P* inclusive 30 

Length of molar series. M^ to M^ ll.S 

Canine — anteroposterior diameter at alveolus: transverse diameter 8.6: 6 

P' — anteroposterior diameter: greatest transverse diameter 8.2: 2.1 

P'^ — anteroposterior diameter: greatest transverse diameter 5.4: 2 

P* — anteroposterior diameter: greatest transverse diameter 8 : 5 

P* — anteroposterior diameter: greatest transverse diameter 10 : 8.2 

M' — anteroposterior diameter: greatest transverse diameter 9.4: 7.5 

M^ — anteroposterior diameter: greatest transverse diameter 3.5: 8.1 

Length of lower jaw from anterior extremity to condyle 92 

Depth of symphysis of lower jaw measured along anterior face 2< 

Depth of flange of lower jaw below diastema between canine and Pi 22 

Depth of lower jaw below Pj 16.6 

Depth of lower jaw below Mj 16.6 

Height of coronoid process above inferior margin of ramus 28 

Length of lower dentition from anterior margin of canine to Mj inclusive 67.6 

Length of cheek tooth series. Pi to M2 inclusive 43.5 

Length of diastema anterior to Pi : posterior to Pi 8 : 2 

Length of premolar series. Pi to P< inclusive 28.8 

Length of molar series. Mi and M2 16.9 

Canine — anteroposterior diameter at cingulum: transverse diameter 6.7: 3.6 

Pi — anteroposterior diameter: greatest transverse diameter 8.6: 1.6 

P2 — anteroposterior diameter: greatest transverse diameter 4.7: 2 

P3 — anteroposterior diameter : greatest transverse diameter 7.6 : 3 

P4 — anteroposterior diameter : greatest transver.se diameter 9.1 : 4 

M, — anteroposterior diameter : greatest transverse diameter 8.6 : 4 

Ma — anteroposterior diameter : greatest transverse diameter 8.7 : 4.4 

Vertebrae. — Among cervical vertebrae preserved of Machae- 



i» Matthew, W. D., ibid., fig. 71. 



MACHAEROroES EOTHEN MATTHEW GAZIN 345 

roides eotheii are the atlas, axis (badly crushed), and three others 
including the seventh. The transverse processes are not complete 
on the atlas, but sufficient remains to show that the groove or 
notch at the anterior extremity for the anterior course of the 
vertebral artery and the inferior branch of the spinal nerve is 
not covered but widely open as in Thinocyon. The posterior open- 
ing of the vertebrarterial foramen, however, is distinctly on the 
posterior border of the transverse process, not dorsal to it as 
described by Matthew for Thinocyon. 

The two intermediate cervicals between the axis and the sev- 
enth show a well-developed inferior lamella with the forward 
ridge well separated from the forward extension of the trans- 
verse process by the opening of the vertebrarterial canal. These 
also show a well-developed forward projecting hyperapophysis 
beginning superior to the postzygapophysis and extending for- 
ward half to three-quarters of the way to articular surface of 
the anterior zygapophysis. It is lower than the spine and more 
compressed transversely. 

The first eight dorsal vertebrae were found in articulation with 
the last cervical, and the centra of seven additional vertebrae, not 
in articulation, were preserved, at least four of which belong to 
the lumbar series. The anterior dorsals show elongate transverse 
processes for articulation with the ribs, and moderately high but 
rapidly tapering spines. The spine of the first dorsal appears to 
be the longest, and possibly the only one to have an anteropos- 
teriorly expanded tip. The centra identified as lumbar are very 
elongate, dorsoventrally flattened, and with the articular faces 
markedly sloping, downward and backward with respect to the 
longitudinal axis. 

Limb bonpfi (pi. 46, c-f) . — The humerus of Machaeroides eothen 
is distinctly larger and more robust than in Limnocyon verus. The 
deltoid and ectocondylar ridges are wide and flaring, and both ex- 
tend for a greater proportion of the length of the shaft than in L. 
veiru?,. The development of the deltoid crest together with the high 
and widely expanded acromion of the .scapula furnisher good lever- 
age and indicates the importance of the deltoid muscU'. which func- 
tions in raising the arm outward. The prominence and length of 
the ectocondylar ridge denote good leverage for the supinator 
longus in flexing and supinating the forearm. The greater and 
lesser tuberosities are well developed with large rugose surfaces 
for the muscles used in rotating the humerus. The distal end of the 
humerus exhibits a large, oval entepicondylar foramen and well- 
developed condyles, particularly the inner, which has a very rugose 



346 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 95 

anterodistal surface for attachment of flexor muscles for the 
manus and forearm. 

The radius and ulna are longer than in Limnocyon verus 
and noticeably curved. Both elements are noticeably flattened 
transversely and anteroposteriorly expanded in their distal por- 
tions, although a part of this is recognized as due to crushing. The 
olecranon of the ulna is relatively robust and bent somev^^hat 
inward but is not so long as in L. verus. The shaft of the ulna is 
not convex anteriorly, as described for L. verus, but, if anything, 
is concave anteriorly. The radius shows a compound curve, con- 
cave forward in the proximal portion and convex forward in the 
medial and distal portions. The proximal portion of the radius 
also has a well-developed bicipital tuberosity, turned slightly 
inward from the shaft of the ulna, for insertion of the biceps. 
Moreover, the distal portion of the radius is very much expanded 
anteromedially above the prominent styloid process. This ex- 
pansion supports the place of insertion for the distal end of the 
supinator longus muscle which had its origin on the well-developed 
supinator ridge of the humerus. 

Remains of the manus include the scapholunar and the third 
and fifth metacarpals. The scaphoid, lunar, and centrale are 
fused, although a groove showing the line of separation between 
the scaphoid and lunar can be seen across a portion of the radial 
facet. The distal facets for articulation with the magnum, unci- 
form, and trapezoid are deeply concave and separated from one 
another by sharp angles. The metacarpals appear short and 
stout, a characteristic noted in machairodonts in comparison with 
true felids. 

Table 2. — Measurevients (in millimeters) of the limb bones of Machaeroides 
eothen, U. S. N. M. No. 17059 

LetiBth of humerus 104 

Anteroposterior diameter of proximal end of humerus ... 26.5 

Transver.se diameter of proximal end of humerus across tuberosities 28 

Transverse diameter of distal end of humerus 81,6 

Greatest length of radius 78,g 

Greatest diamster of proximal end of radius 13.5 

Greatest diameter of distal end of radius ... I8.8 

Lengtli of ulna 102 

Anteroposterior diameter of olecranon below tuberosities 13.6 

Greatest diameter of distal end of ulna 13 

Greatest lenslh of scapholunar Ig 

Length of metacarpal III 27 

Length of metacarpal V 21 

Length of femur parallel to axis of shaft 118 

Diameter of proximal end of femur across head and greater trochanter, perpendicular 

to axis of shaft ^ 35 

Transverse diameter of shaft of femur at midsection 17.5 

Transverse diameter of distal end of femur across condyles, excludine tuberosities 18 

Transverse diameter of disUl end of femur across tuberosities IS 



MACHAEROIDES EOTHEN MATTHEW — GAZIN 347 

Both femora are preserved although the right is partly dam- 
aged. This element is longer and sturdier than in Linmocyon 
vents but otherwise has about the same curvature. The greater 
trochanter, though exhibiting an enlarged, rugose surface, does 
not project upward so far as in L. verus. The lesser trochanter 
is a posterointernal knoblike process situated at about the same 
height on the shaft as in L. verus, but the third trochanter is dis- 
tinctly higher or more proximally located on the shaft, only 
slightly lower than the lesser trochanter. The distd extremity is 
relatively flattened on the dorsal surface and the patellar trochlea 
not so long as in L. verus but with the depression carri'sd upward 
on the shaft well beyond the termination of the articulating sur- 
face. 

The preserved portion of the tibia is relatively robust, and 
curved, as described for L. verus, but the cnemial crest is promi- 
nent and extends distally a greater distance. The proximal ex- 
tremity is missing but the posterior surface shows a rather deep 
groove between the ridges supporting the condyles, emphasized 
in part by crushing. The distal extremity may not be entire. It 
terminates in a simple, nearly flat surface v/ith a steep dorso- 
lateral slope for articulation with the astragalus. The internal 
mTlIcolus may not extend much below the facet for the astragalus, 

t is incomplete. 



^ U. S GOVERNMENT PRINTING OfFlCE ISa-S— 714083 



U. S. NATIONAL MUSEUM 



PROCEEDINGS. VOL 96 PLATE 45 






Mnrharroidc. mtheu Matthew. Skull (U.S.N.M. No. 1T0.V..) : n, Dor.sal view; 
I, iatfial view; r. ventral view; (/. occlusal view of upper cheek teeth 
liiulKi'V Kocene, Wyoniintr. n-c three-fourths natural size; d, I'l: natural 
si/e DrawiiiL' ('/) l>y Mrs. Ainie .M. Awl. 



U. S. NATIONAL MUSEUM 



PROCEEDINGS, VOL. 96 PLATE 46 




-*<*-^«^r 






a 







Machaeroides eothen Matthew, a, b, Mandible (U.S.N.M. No. 17059) : a, 
lingual view of left ramus; b, lateral view of right ramus, c-f, Limb bones 
(U.S.N.M. No. 17059) : c, anterior view of left femur; d, posterior view of 
left humerus; e, medial view of right humerus; /, lateral view of right 
radius and ulna. Bridger Eocene, Wyoming, a, b natural size; c-f three- 
fifths natural size. 




PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 



issued 



SMITHSONIAN INSTITUTION 

U. S. NATIONAL MUSEUM 
Vol. 96 Washington: 1946 No. 3203 

REVIEW OF SOME CHALCIDOID GENERA RELATED TO 
CEROCEPHALA WESTWOOD 



By A. B. Gahan 



This review of some genera related to Cerocephala Westwood 
was occasioned by difficulty experienced in placing satisfactorily 
specimens received for identification. It is hoped that the fol- 
lowing treatment will eliminate some of the existing confusion 
regarding these genera and make identification somewhat more 
certain. 

Cerocephala and its allies have been associated by most authors 
with the genus Spalanf/ia Latreille to form a subfamily Spalangi- 
inae or a tribe Spalangiini in the family Pteromalidae. In my 
opinion Cerocephala and its allies are not closely related to 
Spalanf/ia and should form a separate group for which I propose 
the name Cerocephalinae. Although resembling each other in 
many respects, the two groups have quite different host relations 
and can be separated by several good characters. The Spalangi- 
inae are all parasitic in dipterous puparia, while the Cerocepha- 
linae so far as known are all associated with Coleoptera. 

The Spalangiiiiae and Cerocei)halinae may be separated by the 
following dichotomy: 

Hind tibia with one calcariuni. Antennao inserted at the extreme anterior 
nianfi'i of head, never scpaiated at base by a frontal carina; funicle always 
7-seK-mentefl. Forewing without a callu.s or tuft of bristles at the proximal 
end of the marginal vein and without transverse fuscous band.s; scutcllum 
usually with a punctate cross furrow before apex Spalangiinac Westwood 

Hind tibia with two calcaria. .Antennae in.serted well above extreme ante- 
rior margin of hoad although frequently distinctly below ventral extremities 
of eyes, always separated at base by a frontal carina or prominence; 
funicle 5- or 6-8egmented, oi- rarely 7-segm«Mited in somo males. Forewing 

349 



350 PROCEEDINGS OF THE NATIONAL MUSEUM vol 96 

always with a, callus at proximal end of marginal vein, this callus fre- 
quently but not always bearing a tuft of erect bristles; usually with a 
transverse fuscous band behind stigmal vein and another at juncture of 
submarginal and marginal veins but occasionally without bands; scutellum 
without a cross furrow before apex Cerocephalinae, new subfamily 

Because of their bicalcarate hind tibiae the cerocephalines would 
run to the family Miscogasteridae in Ashmead's key to families 
of Chalcidoidea (Mem. Carnegie Mus., vol. 1, p. 228, 1904) , and in 
that group, as constituted by Ashmead, they closely resemble 
some apterous forms in the subfamily Lelapinae, from which they 
differ principally by the absence of a neck on the propodeum and 
by differences in the shape of the head. I do not consider the 
Miscogasteridae to be separable as a family from the Pteromali- 
dae. Instead, I think it should be combined with the Pteromali- 
dae to form a large family made up of numerous smaller groups 
including the Cerocephalinae, Spalangiinae, and Lelapinae among 
numerous others. 

Family PTEROMALIDAE 
Cerocephalinae, new subfamily 

Eight genera are here included in the Cerocephalinae. viz: 
Choetospila Westwood, Theocolax Westwood, Cerocephala West- 
wood, Theocolaxia Girault, Parcdaesthia Cameron, Acerocephala 
new genus, Neosciatheras Masi, and Sciatherellus Masi. No 
representatives of Paralaesthia, Neosciatheras, and Sciatherellvs 
have been available for study, and they are placed in the generic 
key solely on the basis of the descriptions. No doubt some genera 
have been omitted that eventually may prove to belong here. 

Four genera that have previously been associated by some 
authors with Cerocephala and Spalangia have been excluded from 
Cerocephalinae. These genera are discussed at the end of the 
paper. 

Description. — Head varying in shape from subcircular to dis- 
tinctly oblong; below antennae usually concave, the margins of 
concavity frequently armed with one or more sharp protuberances 
or teeth on each side; if not concave then face with striae con- 
verging toward clypeus ; antennae inserted at or beloM' middle of 
head but always distinctly above clypeus and always separated 
at base by a raised carina or narrow plate which is frequently 
but not always produced anteriorly to form a sharp tooth or spine 
between the antennae; malar groove absent; occiput carinately 
margined above ; head attached to neck of pronotum very close to 
vertex. Pronotum large, subconical, sloping from posterior 
margin to neck without any abrupt angulations either dorsally or 
laterally ; parapsidal grooves distinct and complete ; axillae either 



CHALCIDOIDEA RELATED TO CEROCEPHALA — GAHAN vSl 

just touching at the median line or narrowly separated; scutellum 
nearly flat, without a cross furrow; propodeum not declivous, its 
dorsum lying in a plane only a little lower than that of scutellum, 
without carinae or lateral folds and not produced into a neck 
posteriorly; prepectus moderately large, triangular; mesopleuron 
with a weak femoral impression. Wings either present or absent; 
when winged, marginal vein of forewing as long as submarginal 
or longer, with a distinct callus or thickening at its juncture with 
submarginal, this callus often but not always bearing a tuft of 
erect bristles ; stigmal vein never more than one-fourth as long as 
marginal, usually much shorter; postmarginal vein never longer 
than stigmal, usually weak, and sometimes absent; discal cilia 
absent or vestigial but marginal cilia moderately long and dense; 
hind wing unusually large with the marginellan vein usually 
longer than submarginella. Forewing usually with one or two 
fuscous areas or cross bands. Hind tibia with two slender, 
unequal, and often very short spurs. Abdomen petiolate; oviposi- 
tor usually shortly exserted, rarely half as long as abdomen; 
gaster of female subcylindrical, sometimes flattened above or a 
little compressed from the sides; of male compressed dorsoven- 
trally and subtruncate at apex. Tegument of head and thorax 
usually smooth and polished, occasionally sculptured. 

KEY TO THE KNOWN GENERA OF CEROCEPHALINAE 
j. Funicle 5-segTnented in the female, 6-segmented in the male; club solid 
in both sexes. Forewing with a tuft of erect bristles at junction of 
submarginal and marginal veins (apterous and subapterous individuals 
of both sexes common) 1. Choetospila Westwood 

Funicle 6-segmented in female, 6- or 7-segmented in male; club of female 
either solid or indistinctly 3-segmented; club of male may be solid, 
distinctly 2-8egmented or indistinctly 3-segmented. Foi-ewing with or 
without a tuft of erect bristles at junction of submarginal and marginal 
veins (apterous forms occur) 2 

2. Head, pronotum, mesoscutum, and .scutellum sculptured; axillae obliquely 

sulcate; forewing without a tuft of bristles at junction of marginal 
and submarginal veins 7 

Head dorsally, and mesoscutum entirely smooth and polished; pronotum 
usually smooth but sometimes sculptured; axillae and scutellum usually 
polished but sometimes more or less striated; forewing with or without 
a tuft of bristles 3 

3. Mandibles unusually long and conspicuous 6 
Mandibles normal 4 

4. Antennae inserted very distinctly below a line connecting lower extremi- 

ties of eyes; head viewed anteriorly as long as broad or longer with 
sides parallel; subapterous or fully winged; fully developed wing with 
a tuft of bristles at juncture of submarginal and marginal veins, post- 
marginal vein absent or represented by merely a short stub. 

2. Th<?ocolax Westwood 



352 PROCEEDINGS OF THE NATIONAL MUSEUM voua 

Antennae inserted higher up, never more than slightly below a line con- 
necting lower extremities of the eyes; head viewed anteriorly usually 
narrowing below, rarely with the sides nearly parallel; forewing with 
or without a tuft of bristles at juncture of submarginal and marginal 
veins; postmarginal vein distinct but never longer than stigmal 
vein 5 

5. Foz'ewing with a distinct tuft of bristles at juncture of submarginal 

and marginal veins 3. Cerocephala Westwood 

Forewing with a callus but without a tuft of bristles at juncture of 
submarginal and mai-ginal veins 4. Theocolaxia Girault 

6. Mandibles bidentate at apex; forewing with a tuft of bristles at juncture 

of submarginal and marginal veins . 5. Paralaesthia Cameron 

Mandibles with four distinct teeth at apex; forewing with a callus but 

without a tuft of bristles at juncture of submarginal and marginal 

veins 6. Acerocephala, new genus 

7. Female: Antennae clavate, inserted a little below middle of head; scape 

not extending above vertex; first funicular joint about twice as long as 
pedicel; postmarginal vein subequal to stigmal vein, the latter rather 
short, the stigmal knob armed with a short tooth. 

7. Neosciatheras Masi 
Male: Antennae very long, filiform, inserted a little above middle of 
head; scape extending above vertex; funicular segments and club 
elongate, cylindrical; pedicel short; postmarginal vein nearly effaced; 
stigmal vein long, straight, and with the stigmal knob scarcely devel- 
oped and unarmed 8. Sciatherellus Masi 

1. Genus CHOETOSPILA Westwood 

Choetospila Westwood, Thesaurus entomologicus Oxoniensis, p, 137, 1874. 
Choetospila Dalla Torre, Catalogus hymenopterorum, vol. 5, p. 207, 1898. 
Spalmigiomor'pha GiRAULT, Mem. Queensland Mus., vol, 2, p. 333, 1913. 

Choetospila has been treated as a synonym of Cerocephala 
Westwood by Ashmead (Mem. Carnegie Mus., vol. 1, p. 369, 1904) 
and others but may be easily distinguished from that genus by 
the 5-segmented funicle of the female. 

Spalangiomorpha was synonymized with Choetospila by Girault 
(Insecutor Inscitiae Menstruus, vol. 5, p. 37, 1917) and later with 
Cerocephala by the same author in one of his privately published 
pamphlets [Indications (in new insects) of ruling power in 
nature, p. 3, 1925]. In a still later publication Girault (Trans, 
and Proc. Roy Soc. South Australia, vol. 53, p. 319. 1929) again 
referred to the genotype species, using the generic name Spalan- 
giomorpha but pointing out that the species agreed with Choeto- 
spila as characterized in a table of genera by Masi (Nov. Zool., 
vol. 24, p. 188, 1917). I agree with Waterston (Rep. Grain Pest 
Committee. No. 9, p. 25, fig. 13, 1921) that Spalangiomorpha is 
certainly a synonym of Choetospila Westwood. 

Tiipe of the genus. — Choetospila elegans Westwood. 



CHALCIDOIDEA RELATED TO CEROCEPHALA — GAHAN 353 

CHOhTOSPlLA ELEGANS Wfstwood 

Plate 47, Figures 1, la 
Choetoapila elegaus Westwood, Thesaurus entomologicus Oxoniensis, p. 157, 

pi. 25, fig. 10, 1874.-- Waterston, Rep. Grain Pe.«t Committee, No. 9, p. 

25, fig. 13, 1921. 
Spalangioinoi pha fasciativentris fJlRAULT, Mem. Queensland Mus., vol. 2, p. 

334, 1913. 
Spalaiigin metallica Fullaway, Proc. Hawaiian Ent. Soc, vol. 2, p. 286, 

1013. (New synonjTny.) 

Spalangiomorpha fasciativentris Girault is certainly the same 
as Choetospila elegans, as was pointed out by Waterston in the 
report cited above. Both the description and the habitat lead to 
this conchision. 

The type of Spalangia metallica Fullaway is in the collection of 
the U. S. National Museum, and it differs in no way from typical 
(\ ilcgaus. 

Choetospila elegans is parasitic upon stored grain weevils and 
has been distributed to many parts of the world in shipments of 
stored products. The National Museum collection contains speci- 
mens from various parts of the United States, Mexico, Puerto 
Rico. Panama Canal Zone, Surinam, Peru, Hawaii. Guam, South 
Australia, Java, India, Nigeria, and South Africa. Published 
records include many additional areas where the species occurs. 

Very little seems to be known about the actual biolog>^ of C. 
elegans beyond the fact that it is usually associated with pests of 
stored ^ain. Its most common host apparently is the rice weevil. 
Sitophihi.'^ orifza (Linnaeus). 5?. grdnaria (Linnaeus). 5>. linearis 
Herbst, Sitodrepa panicea (Linnaeus), Caulophilus latinasus Say, 
Callofioh'nichus quadrinmcidatns (Fabricius), and C. chinensis 
(Linnaeus) have been recorded as attacked by it. Very likely 
other grain-infesting beetles may serve as hosts, but no authentic 
records of such hosts have been published. A paper by Corbett 
and Miller (Federated Malay States Dept. Agr. Sci.. ser. 1P>, p. 4. 
H).3.''>) recording this species as a parasite of Sitotroga cerealella 
Olivier is not available for judgment as to the authenticity of the 
record. Squire (T)iel Rep. Dept. Agri. British Guiana for 1924, 
pp. 121 124, 1925) records C\ rhftnns as parasitizing an uniden- 
tiled scolytid infesting British Honduras mahogany tn^es (Sirri- 
frnia /??/?«?7i,s-) in Guiana, and in the U. S. National Museum 
collection are 14 specimens of C. rlninyix labeled as having been 
reared by J. Zetek in 1924 in the l^anama Ganal Zone from a 
scolytid-infested piece of "Amargo" wood iVntnirra sp.). 

The species is sufficiently characterized by the descriptions 
and figures given in the above-cited papers by Westwood and 
Waterston. 



354 PROCEEDINGS OF THE NATIONAL MUSEUM VOL 96 

CHOETOSPILA PRATER (Girault), new combination 

Spalangiomorpha frater GiRAULT, Mem. Queensland Mus., vol. 2, p. 334, 1913. 

No representative of this Australian species is available for 
study. According to the description it differs from elegans prin- 
cipally by having the first funicular joint wider than long. 

CHOETOSPILA TABIDA, new species 

Plate 47, Figures 2, 2a 

This species is very similar to elegans but differs by having 
distinctly shorter antennae, by being entirely wingless in both 
sexes (so far as shown by the specimens at hand) , by the propo- 
deum being smooth and polished, by the ovipositor of the female 
not extending beyond the apex of the abdomen, and by the an- 
tennal club being unsegmented in both sexes. 

Female. — Length 1,6 mm. Mostly smooth and polished, the 
head anteriorly, below level of insertion of antennae, weakly 
wrinkled, and the mesopleura reticulated. 

Head viewed from in front a little longer than broad, nearly 
squarely truncate at vertex and at mouth, and with the sides 
practically parallel; compound eyes very small, subobsolete; 
ocelli represented merely by minute punctures ; antennae inserted 
low down at approximately the lower fifth of head, separated at 
base by a sharp carina ; scrobes short and shallow, poorly defined ; 
face slightly depressed medially, with a small toothlike projection 
at the anterior margin on each side of clypeus ; mandibles rather 
broad and stout. Head viewed laterally forming a rather broad 
ellipse, approximately twice as long as broad. 

Antenna 8-segmented, strongly clavate, short, its total length 
only slightly greater than the frontal length of head ; scape short, 
slightly thickened, about three times as long as broad; pedicel 
about half as long as scape and half as long as funicle; funicle 
5-segmented, the segments all transverse and gradually increasing 
in width toward club; club ovoid, large, very nearly as long as 
entire funicle, approximately twice as long as broad and appar- 
ently not segmented. 

Thorax a little narrower than head, approximately twice as long 
as broad; prothorax subconical, rounded at the sides, more than 
twice as long as mesoscutum; parapsidal grooves complete, very 
oblique; scutellum as long as mesoscutum, nearly circular; meso- 
pleuron weakly reticulated; metapleuron polished; propodeum 
slightly longer than scutellum, weakly convex dorsally, without 
grooves or carinae, smooth and polished except for some very 
faint reticulation basad of the spiracles. Wings entirely absent. 

Legs rather stout, the middle pair obviously somewhat shorter 



CHALCIDOIDEA RELATED TO CEROCEPHALA — GAHAN ^>S 

and weaker than the other two pairs; anterior femur obviously 
.swollen and its tibia a little thickened, each with a few moderately 
longr hairs on its ventral margin and a few shorter ones on its 
(^uter face; posterior femur slijjrhtly swollen, with a few short 
hairs; posterior tibia slender at base but gradually increasing in 
thickness toward apex, sparsely hairy and with two rather long- 
and slender but unequal calcaria ; tarsi not thickened ; posterior 
basitarsus equal to half the length of tibia. 

Abdomen nearly as long as head and thorax combined, a little 
broader than thorax ; petiole as long as broad, smooth ; gaster 
elliptical in outline, twice as long as broad, smooth and polished 
dorsally and ventrally, nearly devoid of hairs dorsally but with 
numerous rather long slender bristles at apex beneath ; first seg- 
ment of gaster emarginate medially: ovipositor not protruding 
Ijeyond apex of abdomen. 

Head, thorax, legs, and abdominal petiole brownish yellow; 
anterior and posterior coxae whitish except dorsally; antennae 
concolorous with head except that club is usually, thouph not 
always, blackish; gaster wholly black or blackish. 

Male. — Length 0.9 mm. Antennae uniformly testaceous, 9- 
.segmented ; funicle 6-segmented. the first four segments very short 
and difficult to distinguish, the last two larger; club solid, conical, 
a little longer than two preceding segments combined; abdomen 
about as long as thorax. Otherwise like the female and difficult 
10 distinguish from it. 

Type locality. — University Park, Md. 

Type.—U. S. N. M. No. 57279. 

fiemarks. — Described from five females (one holotype) and one 
male collected by W. H. Anderson as pupae from the burrows of 
an unidentified cossonine beetle infesting a dead branch of an 
unidentified tree and reared to adults. 

2. Genus THEOCOLAX Wcstwood 

'! hfocalax Westwood, Philos. Map., ser. .3, vol. 1. p. ll!7. IhV'. 
Lnrythia Halihay, Ent. .Map., vol. 1, p. 3:55, 18.3.3. 

Theocolax is very similar to Crrocephala, and it has been treated 
as a synonym by Walker (Ent. Mag., vol. 2, p. 148, 1833). It 
may be distinguishod from Cmirrphahi, howt'vor. l\v h;;ving the 
antennae inserted distinctly below a line connecting the lower 
extremities of the eye.s, the head viewed anteriorly as long as 
broad or a little lon^rer, parallel-sided, and never with more than 
three small toothlike projections, one between the antennae and 
one on each side of the facial depression just above the base of 
each mandible. In some species the processes above the bases 



356 PROCEEDINGS OF THE NATIONAL MUSEUM VOL 96 

of the mandibles are absent. Both subapterous and fully winged 
individuals occur. In the fully winged individuals there is a 
distinct tuft of erect hairs at the juncture of the marginal and 
submarginal veins of the forewing, exactly as in Cerocephala. 
The postmarginal vein is absent or represented by merely a short 
stub. 

Type of the genus. — Theocolax foiiniciformis Westwood. 

THEOCOLAX FORMICIFORMIS Westwood 

Plate 47, Figs. 3, 3a; Plate 48, Figs. 1, la, lb 

Theocolax formiciformis Westwood, Philos. Mag., ser. 3, vol. 1, p. 127, 1832; 

Thesaurus entomologicus Oxoniensis, p. 138, pi. 25, fig. 11, 1874. 
Laesthia vespertina Haliday, Ent. Mag., vol. 1, p. 336, 1833. 
Cerocephala formiciformis (Westwood) Walker, Ent. Mag., vol. 2, p. 149, 

1834. 

I examined Westwood's type of this species in the Hope 
Museum at Oxford, England, in 1927 and compared with it a 
specimen from Blankenburg, Thuringia, previously identified by 
Schmiedeknecht. This specimen, now in the U. S. National Mu- 
seum, appeared to me to be homotypic. The National Museum 
collection also contains one specimen identified by Ruschka and 
said to have been reared from Anobium striatum Olivier at Stock- 
holm, Sweden. This specimen is considerably paler in color than 
the homotype but does not seem to differ otherwise. In addition 
to these two specimens the collection possesses a series of 14 
specimens from Auckland, New Zealand, said to have been reared 
from Anobium-mf ested timber by E. Bollard and D. Spiller. 
Apparently this series does not differ materially from the homo- 
type except that one female in the lot has fully developed wings. 
Except for the perfectly developed wings, this winged specimen 
is exactly like the subapterous females. 

Although the species has usually been described as wingless, as 
a matter of fact, in none of the individuals I have seen are the 
wings completely absent, usually being represented by small stubs 
approximately twice the length of the tegula. In the fully winged 
individual the postmarginal vein is represented by merely a very 
short stub ; the marginal vein is a little longer than the submar- 
ginal; the stigmal vein is approximately one-sixth as long as 
marginal, slightly curved and not thickened at apex; the disk of 
wing is nearly bare ; the marginal cilia are moderately long, and 
there is an erect tuft of black bristles at the junction of marginal 
and submarginal veins. There is a broad fuscous cloud embracing 
the apical half of the marginal and all of the stigmal vein and 
extending across the wing. The hind wing is about two-thirds as 



CHALCIDOIDEA RELATED TO CEROCEPHALA — GAHAN 357 

wide as the forewing and rounded at its apex. The propodeum is 
weakly reticulated and slightly shining. 

Theocolax formiciformis appears to be a common parasite of 
Anohium. having been recorded from several species of that 
genus. It is also said by several authors to attack (Hylesinus) 
Leperisinus fraxini (Panzer), but since it is known to have been 
confused in some instances with Cerocephala corniyera Westwood. 
a common parasite of this host, it seems very likely that all of 
the records of Theocolax formiciformifi from Leperisiuus fraxini 
actually refer to Cerocephala cornigea as was suggested by 
Waterston (Rep. Grain Pests Committee, No. 9. p. 12. 1921). 

The species is known to occur in England, Sweden. Germany, 
and New Zealand and may be much more widely distributed. It 
is not known to occur in America. 

THEOCOLAX BAKERI (Crawford), new combination 

'^rrocephnla bakfri CRAWFORD. Philippine Journ. Sci., vol. 0, p. 4t:10. 1914. 

This Philippine species differs from formiciformis by being 
distinctly smaller, by lacking the toothlike protuberances on each 
side of the facial depression, and by the pedicel in the female 
being as long as the first and second segments of the funicle com- 
bined. It is also somewhat paler in color. All known specimens 
of the species are fully winged and have a conspicuous tuft of 
bristles at the .junction of the marginal and submarginal veins. 
The head is oblong with parallel sides and with the antennae 
inserted distinctly below the ventral extremities of the eyes. 

There are eight specimens in the U. S. National Museum, 
inclusive of the type series, all taken by C. F. Baker at Los Banos, 
Philippine Islands. One of these bears a label "on scolytid No. 
847." 

THEOCOLAX LITIGIOSA (Kondani) 

Laeathia liiigiosa Rondani, Ann. Soc. Nat. Modena, vol. 1, p. 23. 1866: Arch. 

Zool., vol. 4, p. 191, pi. 7, fijf. 7-8, 1866; Bull. Soc. Ent. Ital., vol. 9, p. 183, 

pi. 1, fig. 20, 1877. 
Theocolax litigioHn Dalla Torre, Catalogus Hymenopteroi iim. vol. 5